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Egg incubation is the process by which an
egg An egg is an organic vessel grown by an animal to carry a possibly fertilized egg cell (a zygote) and to incubate from it an embryo within the egg until the embryo has become an animal fetus that can survive on its own, at which point the ...
, of
oviparous Oviparous animals are animals that reproduce by depositing fertilized zygotes outside the body (i.e., by laying or spawning) in metabolically independent incubation organs known as eggs, which nurture the embryo into moving offsprings kno ...
(egg-laying) animals, develops an
embryo An embryo ( ) is the initial stage of development for a multicellular organism. In organisms that reproduce sexually, embryonic development is the part of the life cycle that begins just after fertilization of the female egg cell by the male sp ...
within the egg, after the egg's formation and
oviposition The ovipositor is a tube-like organ used by some animals, especially insects, for the laying of eggs. In insects, an ovipositor consists of a maximum of three pairs of appendages. The details and morphology of the ovipositor vary, but typica ...
al release. Egg incubation is done under favorable environmental conditions, possibly by brooding and hatching the egg. Multiple and various factors are vital to the incubation of various species of animal. In many species of reptile for example, no fixed temperature is necessary, but the actual temperature determines the sex ratio of the offspring. In birds, the sex of offspring is genetically determined, but in many species a constant and particular
temperature Temperature is a physical quantity that quantitatively expresses the attribute of hotness or coldness. Temperature is measurement, measured with a thermometer. It reflects the average kinetic energy of the vibrating and colliding atoms making ...
is necessary for successful incubation. Especially in
poultry Poultry () are domesticated birds kept by humans for the purpose of harvesting animal products such as meat, Eggs as food, eggs or feathers. The practice of animal husbandry, raising poultry is known as poultry farming. These birds are most typ ...
, the act of sitting on eggs to incubate them is called brooding. The action or behavioral tendency to sit on a clutch of eggs is also called
broodiness Broodiness is the action or behavioral tendency to sit on a clutch of eggs to incubate them, often requiring the non-expression of many other behaviors including feeding and drinking.Homedes Ranquini, J. y Haro-García, F. Zoogenética. 1ra. edi ...
, and most
egg An egg is an organic vessel grown by an animal to carry a possibly fertilized egg cell (a zygote) and to incubate from it an embryo within the egg until the embryo has become an animal fetus that can survive on its own, at which point the ...
-laying breeds of poultry have had this behavior
selectively bred Selective breeding (also called artificial selection) is the process by which humans use animal breeding and plant breeding to selectively develop particular phenotypic traits (characteristics) by choosing which typically animal or plant ma ...
out of them to increase production.


Avian incubation

A wide range of incubation habits is displayed among birds. In
warm-blooded Warm-blooded is a term referring to animal species whose bodies maintain a temperature higher than that of their environment. In particular, homeothermic species (including birds and mammals) maintain a stable body temperature by regulating ...
species such as
bird Birds are a group of warm-blooded vertebrates constituting the class (biology), class Aves (), characterised by feathers, toothless beaked jaws, the Oviparity, laying of Eggshell, hard-shelled eggs, a high Metabolism, metabolic rate, a fou ...
species generally, body heat from the
brooding patch A brood patch, also known as an incubation patch in older literature, is a patch of featherless skin on the underside of birds during the nesting season. Feathers act as inherent insulators and prevent efficient incubation, to which brood patches ...
of the brooding parent provides the constant temperature. Several groups, notably the
megapode The megapodes, also known as incubator birds or mound-builders, are stocky, medium-large, chicken-like birds with small heads and large feet in the family Megapodiidae. Their name literally means "large foot" and is a reference to the heavy leg ...
s, instead use heat generated from rotting vegetable material, effectively creating a giant
compost Compost is a mixture of ingredients used as plant fertilizer and to improve soil's physical, chemical, and biological properties. It is commonly prepared by Decomposition, decomposing plant and food waste, recycling organic materials, and man ...
heap, while
crab plover The crab-plover or crab plover (''Dromas ardeola)'' is a coastal wader (shorebird). It is the only member of the genus ''Dromas'' and the family Dromadidae. It is unique among waders in making use of ground warmth to aid the incubation of its e ...
s make partial use of heat from the sun. The
Namaqua sandgrouse The Namaqua sandgrouse (''Pterocles namaqua''), is a species of ground-dwelling bird in the sandgrouse family. It is found in arid regions of south-western Africa. Taxonomy The Namaqua sandgrouse was formally described in 1789 by the German na ...
of the deserts of southern
Africa Africa is the world's second-largest and second-most populous continent after Asia. At about 30.3 million km2 (11.7 million square miles) including adjacent islands, it covers 20% of Earth's land area and 6% of its total surfac ...
, needing to keep its eggs cool during the heat of the day, stands over them drooping its wings to shade them. The humidity is also critical, because if the air is too dry the egg will lose too much water to the atmosphere, which can make hatching difficult or impossible. As incubation proceeds, an egg will normally become lighter, and the air space within the egg will normally become larger, owing to evaporation from the egg. During incubation, the inner layers of the shell are dissolved by their acidic environment and the calcium carbonate that had been part of the shell is incorporated into the skeleton of the foetus. Experiments with
great tit The great tit (''Parus major'') is a small passerine bird in the tit family Paridae. It is a widespread and common species throughout Europe, the Middle East, Central Asia and east across the Palearctic to the Amur River, south to parts of No ...
s show that females compensate for the potential effects of differential heating by moving the eggs homogeneously within the clutch. In the species that incubate, the work is divided differently between the sexes. Possibly the most common pattern is that the female does all the incubation, as in the
Atlantic canary The Atlantic canary (''Serinus canaria''), known worldwide simply as the wild canary and also called the island canary, common canary, or canary, is a small passerine bird belonging to the genus '' Serinus'' in the true finch family, Fringillid ...
and the
Indian robin The Indian robin (''Copsychus fulicatus'')Rasmussen & Anderton emend the species epithet from ''fulicata'' to ''fulicatus'' since ''Saxicola'' is masculine and the ''-oides'' ending is always masculine according to ICZN Code 30.1.4.4ICZN Code. Se ...
, or most of it, as is typical of
falcon Falcons () are birds of prey in the genus ''Falco'', which includes about 40 species. Some small species of falcons with long, narrow wings are called hobbies, and some that hover while hunting are called kestrels. Falcons are widely distrib ...
s. In some species, such as the
whooping crane The whooping crane (''Grus americana'') is an endangered Crane (bird), crane species, native to North America, named for its "whooping" calls. Along with the sandhill crane (''Antigone canadensis''), it is one of only two crane species native to ...
, the male and the female take turns incubating the egg. In others, such as the
cassowaries Cassowaries (; Biak language, Biak: ''man suar'' ; ; Papuan_languages, Papuan: ''kasu weri'' ) are flightless birds of the genus ''Casuarius'', in the order Casuariiformes. They are classified as ratites, flightless birds without a keel (bird a ...
, only the male incubates. The male
mountain plover The mountain plover (''Anarhynchus montanus'') is a medium-sized ground bird in the plover family ( Charadriidae). It is misnamed, as it lives on level land. Unlike most plovers, it is usually not found near bodies of water or even on wet soil; ...
incubates the female's first clutch, but if she lays a second, she incubates it herself. In
hoatzin The hoatzin ( ) or hoactzin ( ) (''Opisthocomus hoazin'') is a species of tropical bird found in swamps, riparian forests, and mangroves of the Amazon and the Orinoco basins in South America. It is the only extant species in the genus ''Opisthoco ...
s, some birds (mostly males) help their parents incubate later broods. The incubation period, the time from the start of uninterrupted incubation to the emergence of the young, varies from 11 days (some small
passerine A passerine () is any bird of the order Passeriformes (; from Latin 'sparrow' and '-shaped') which includes more than half of all bird species. Sometimes known as perching birds, passerines generally have an anisodactyl arrangement of their ...
s and the black-billed and
yellow-billed cuckoo The yellow-billed cuckoo (''Coccyzus americanus'') is a member of the cuckoo family. Common folk names for this bird in the southern United States are rain crow and storm crow. These likely refer to the bird's habit of calling on hot days, often ...
s) to 85 days (the
wandering albatross The snowy albatross (''Diomedea exulans''), also known as the wandering albatross, white-winged albatross, or goonie, is a large seabird from the family Diomedeidae Albatrosses, of the biological family (biology), family Diomedeidae, are la ...
and the brown kiwi). In these latter, the incubation is interrupted; the longest uninterrupted period is 64 to 67 days in the
emperor penguin The emperor penguin (''Aptenodytes forsteri'') is the tallest and heaviest of all living penguin species and is Endemism in birds, endemic to Antarctica. The male and female are similar in plumage and size, reaching in length and weighing fr ...
. In general smaller birds tend to hatch faster, but there are exceptions, and cavity nesting birds tend to have longer incubation periods. It can be an energetically demanding process, with adult albatrosses losing as much as 83 g of body weight a day.
Megapode The megapodes, also known as incubator birds or mound-builders, are stocky, medium-large, chicken-like birds with small heads and large feet in the family Megapodiidae. Their name literally means "large foot" and is a reference to the heavy leg ...
eggs take from 49 to 90 days depending on the mound and ambient temperature. Even in other birds, ambient temperatures can lead to variation in incubation period. Normally the egg is incubated outside the body. However, in one recorded case, the egg incubation occurred entirely within a chicken. The chick hatched inside and emerged from its mother without the shell, leading to internal wounds that killed the mother hen. Embryo development remains suspended until the onset of incubation. The freshly laid eggs of domestic fowl, ostrich, and several other species can be stored for about two weeks when maintained under 5 °C. Extended periods of suspension have been observed in some marine birds. Some species begin incubation with the first egg, causing the young to hatch at different times; others begin after laying the second egg, so that the third chick will be smaller and more vulnerable to food shortages. Some start to incubate after the last egg of the clutch, causing the young to hatch simultaneously. Incubation periods for birds:


Mammalian incubation

The only living
mammal A mammal () is a vertebrate animal of the Class (biology), class Mammalia (). Mammals are characterised by the presence of milk-producing mammary glands for feeding their young, a broad neocortex region of the brain, fur or hair, and three ...
s that lay eggs are
echidna Echidnas (), sometimes known as spiny anteaters, are quill-covered monotremes (egg-laying mammals) belonging to the Family (biology), family Tachyglossidae , living in Australia and New Guinea. The four Extant taxon, extant species of echidnas ...
s and
platypus The platypus (''Ornithorhynchus anatinus''), sometimes referred to as the duck-billed platypus, is a semiaquatic, egg-laying mammal endemic to eastern Australia, including Tasmania. The platypus is the sole living representative or monotypi ...
es. In the latter, the eggs develop ''
in utero The uterus (from Latin ''uterus'', : uteri or uteruses) or womb () is the organ in the reproductive system of most female mammals, including humans, that accommodates the embryonic and fetal development of one or more fertilized eggs until bir ...
'' for about 28 days, with only about 10 days of external incubation (in contrast to a chicken egg, which spends about one day in tract and 21 days externally). After laying her eggs, the female curls around them. The incubation period is divided into three phases. In the first phase, the
embryo An embryo ( ) is the initial stage of development for a multicellular organism. In organisms that reproduce sexually, embryonic development is the part of the life cycle that begins just after fertilization of the female egg cell by the male sp ...
has no functional organs and relies on the
yolk sac The yolk sac is a membranous wikt:sac, sac attached to an embryo, formed by cells of the hypoblast layer of the bilaminar embryonic disc. This is alternatively called the umbilical vesicle by the Terminologia Embryologica (TE), though ''yolk sac' ...
for sustenance. The yolk is absorbed by the developing young. During the second phase, the digits develop. In the last phase, the
egg tooth An egg tooth is a temporary, sharp projection present on the bill or snout of an oviparous animal at hatching. It allows the hatchling to penetrate the eggshell from inside and break free. Birds, reptiles, and monotremes possess egg teeth as h ...
appears.


Reptilian incubation

Methods of incubation vary widely among the many different kinds of reptiles. Various species of
sea turtle Sea turtles (superfamily Chelonioidea), sometimes called marine turtles, are reptiles of the order Testudines and of the suborder Cryptodira. The seven existing species of sea turtles are the flatback, green, hawksbill, leatherback, loggerh ...
s bury their eggs on beaches under a layer of sand that provides both protection from predators and a constant temperature for the nest.
Snake Snakes are elongated limbless reptiles of the suborder Serpentes (). Cladistically squamates, snakes are ectothermic, amniote vertebrates covered in overlapping scales much like other members of the group. Many species of snakes have s ...
s may lay eggs in communal burrows, where a large number of adults combine to keep the eggs warm. Some species coil their torsos around the eggs to provide heat for incubation.
Alligator An alligator, or colloquially gator, is a large reptile in the genus ''Alligator'' of the Family (biology), family Alligatoridae in the Order (biology), order Crocodilia. The two Extant taxon, extant species are the American alligator (''A. mis ...
s and
crocodile Crocodiles (family (biology), family Crocodylidae) or true crocodiles are large, semiaquatic reptiles that live throughout the tropics in Africa, Asia, the Americas and Australia. The term "crocodile" is sometimes used more loosely to include ...
s either lay their eggs in mounds of decomposing vegetation or lay them in holes they dig in the ground.


Incubation by other vertebrates

Fish generally do not incubate their eggs. However, some species
mouthbrood Mouthbrooding, also known as oral incubation and buccal incubation, is the care given by some groups of animals to their offspring by holding them in the mouth of the parent for extended periods of time. Although mouthbrooding is performed by a va ...
their eggs, not eating until they hatch. Some amphibians brood their eggs. The female salamander ''
Ensatina The ensatina (''Ensatina eschscholtzii'') is a species complex of plethodontid (lungless) salamanders found in coniferous forests, oak woodland and chaparral
'' (''Ensatina eschscholtzii'') curls around the clutch of eggs and massages individual eggs with her pulsating throat. Some aquatic frogs such as the Surinam toad (''
Pipa pipa The pipa, pípá, or p'i-p'a () is a traditional Chinese musical instrument belonging to the plucked category of instruments. Sometimes called the "Chinese lute", the instrument has a pear-shaped wooden body with a varying number of frets rangi ...
'') have pouches in their skin into which the eggs are inserted. Other neotropical frogs in the family
Hemiphractidae The Hemiphractidae are a family of frogs from South and Central America. Previously, this group had been classified as a subfamily (Hemiphractinae) under family Hylidae. More recent research classifies these genera into their own family, or som ...
also have pouches in which the eggs develop, in some species directly into juvenile frogs and in others into tadpoles that are later deposited in small water bodies to continue their development. The male
Darwin's frog Darwin's frog (''Rhinoderma darwinii''), also called the Southern Darwin's frog, is a species of frog of the family Rhinodermatidae. It was discovered by Charles Darwin during his voyage on HMS ''Beagle''. on a trip to Chile. In 1841, French ...
carries the eggs around in his mouth until
metamorphosis Metamorphosis is a biological process by which an animal physically develops including birth transformation or hatching, involving a conspicuous and relatively abrupt change in the animal's body structure through cell growth and different ...
, and the female stomach-brooding frog of Australia swallows the eggs, which develop in her stomach.


Incubation by invertebrates

Brooding occurs in some invertebrates when the fertilised eggs are retained inside or on the surface of the parent, usually the mother. This happens in some
cnidaria Cnidaria ( ) is a phylum under kingdom Animalia containing over 11,000 species of aquatic invertebrates found both in fresh water, freshwater and marine environments (predominantly the latter), including jellyfish, hydroid (zoology), hydroids, ...
ns (
sea anemone Sea anemones ( ) are a group of predation, predatory marine invertebrates constituting the order (biology), order Actiniaria. Because of their colourful appearance, they are named after the ''Anemone'', a terrestrial flowering plant. Sea anemone ...
s and
coral Corals are colonial marine invertebrates within the subphylum Anthozoa of the phylum Cnidaria. They typically form compact Colony (biology), colonies of many identical individual polyp (zoology), polyps. Coral species include the important Coral ...
s), a few
chiton Chitons () are marine molluscs of varying size in the class Polyplacophora ( ), formerly known as Amphineura. About 940 extant and 430 fossil species are recognized. They are also sometimes known as sea cradles or coat-of-mail shells or suck ...
s, some gastropod molluscs, some
cephalopod A cephalopod is any member of the molluscan Taxonomic rank, class Cephalopoda (Greek language, Greek plural , ; "head-feet") such as a squid, octopus, cuttlefish, or nautilus. These exclusively marine animals are characterized by bilateral symm ...
s, some
bivalve molluscs Bivalvia () or bivalves, in previous centuries referred to as the Lamellibranchiata and Pelecypoda, is a class of aquatic molluscs (marine and freshwater) that have laterally compressed soft bodies enclosed by a calcified exoskeleton consis ...
, many
arthropod Arthropods ( ) are invertebrates in the phylum Arthropoda. They possess an arthropod exoskeleton, exoskeleton with a cuticle made of chitin, often Mineralization (biology), mineralised with calcium carbonate, a body with differentiated (Metam ...
s, some
entoprocta Entoprocta (), or Kamptozoa , is a phylum (biology), phylum of mostly Sessility (zoology), sessile aquatic animals, ranging from long. Mature individuals are goblet-shaped, on relatively long stalks. They have a "crown" of solid tentacles whos ...
ns, some
brachiopod Brachiopods (), phylum (biology), phylum Brachiopoda, are a phylum of animals that have hard "valves" (shells) on the upper and lower surfaces, unlike the left and right arrangement in bivalve molluscs. Brachiopod valves are hinged at the rear e ...
s, some
bryozoa Bryozoa (also known as the Polyzoa, Ectoprocta or commonly as moss animals) are a phylum of simple, aquatic animal, aquatic invertebrate animals, nearly all living in sedentary Colony (biology), colonies. Typically about long, they have a spe ...
ns, and some
starfish Starfish or sea stars are Star polygon, star-shaped echinoderms belonging to the class (biology), class Asteroidea (). Common usage frequently finds these names being also applied to brittle star, ophiuroids, which are correctly referred to ...
.


See also

* Artificial incubation *
Broodiness Broodiness is the action or behavioral tendency to sit on a clutch of eggs to incubate them, often requiring the non-expression of many other behaviors including feeding and drinking.Homedes Ranquini, J. y Haro-García, F. Zoogenética. 1ra. edi ...
*
Incubator (egg) An incubator is a device simulating avian incubation by keeping egg (biology), eggs warm at a particular temperature range and in the correct humidity with a turning mechanism to hatch them. The common names of the incubator in other terms inclu ...


References

*


Further reading

*


External links


Incubation periods in table
from PennState Extension {{Authority control Bird breeding Eggs Reproduction in animals Ethology sv:Häckning#Ruvning