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Metamorphosis is a
biological process Biological processes are those processes that are vital for an organism to live, and that shape its capacities for interacting with its environment. Biological processes are made of many chemical reactions or other events that are involved in the ...
by which an animal physically develops including birth or hatching, involving a conspicuous and relatively abrupt change in the animal's body structure through cell
growth Growth may refer to: Biology * Auxology, the study of all aspects of human physical growth * Bacterial growth * Cell growth * Growth hormone, a peptide hormone that stimulates growth * Human development (biology) * Plant growth * Secondary grow ...
and differentiation. Some
insect Insects (from Latin ') are pancrustacean hexapod invertebrates of the class Insecta. They are the largest group within the arthropod phylum. Insects have a chitinous exoskeleton, a three-part body ( head, thorax and abdomen), three pa ...
s,
fish Fish are Aquatic animal, aquatic, craniate, gill-bearing animals that lack Limb (anatomy), limbs with Digit (anatomy), digits. Included in this definition are the living hagfish, lampreys, and Chondrichthyes, cartilaginous and bony fish as we ...
,
amphibian Amphibians are four-limbed and ectothermic vertebrates of the class Amphibia. All living amphibians belong to the group Lissamphibia. They inhabit a wide variety of habitats, with most species living within terrestrial, fossorial, arbo ...
s,
mollusk Mollusca is the second-largest phylum of invertebrate animals after the Arthropoda, the members of which are known as molluscs or mollusks (). Around 85,000  extant species of molluscs are recognized. The number of fossil species is e ...
s,
crustacean Crustaceans (Crustacea, ) form a large, diverse arthropod taxon which includes such animals as decapoda, decapods, ostracoda, seed shrimp, branchiopoda, branchiopods, argulidae, fish lice, krill, remipedes, isopoda, isopods, barnacles, copepods, ...
s,
cnidaria Cnidaria () is a phylum under kingdom Animalia containing over 11,000 species of aquatic animals found both in freshwater and marine environments, predominantly the latter. Their distinguishing feature is cnidocytes, specialized cells that ...
ns,
echinoderm An echinoderm () is any member of the phylum Echinodermata (). The adults are recognisable by their (usually five-point) radial symmetry, and include starfish, brittle stars, sea urchins, sand dollars, and sea cucumbers, as well as the ...
s, and
tunicate A tunicate is a marine invertebrate animal, a member of the subphylum Tunicata (). It is part of the Chordata, a phylum which includes all animals with dorsal nerve cords and notochords (including vertebrates). The subphylum was at one t ...
s undergo metamorphosis, which is often accompanied by a change of
nutrition Nutrition is the biochemical and physiological process by which an organism uses food to support its life. It provides organisms with nutrients, which can be metabolized to create energy and chemical structures. Failure to obtain sufficient ...
source or behavior. Animals can be divided into species that undergo complete metamorphosis (" holometaboly"), incomplete metamorphosis (" hemimetaboly"), or no metamorphosis (" ametaboly"). Scientific usage of the term is technically precise, and it is not applied to general aspects of
cell growth Cell growth refers to an increase in the total mass of a cell, including both cytoplasmic, nuclear and organelle volume. Cell growth occurs when the overall rate of cellular biosynthesis (production of biomolecules or anabolism) is greater th ...
, including rapid growth spurts. Generally organisms with a
larva A larva (; plural larvae ) is a distinct juvenile form many animals undergo before metamorphosis into adults. Animals with indirect development such as insects, amphibians, or cnidarians typically have a larval phase of their life cycle. ...
stage undergo metamorphosis, and during metamorphosis the organism loses larval characteristics. References to "metamorphosis" in
mammal Mammals () are a group of vertebrate animals constituting the class Mammalia (), characterized by the presence of mammary glands which in females produce milk for feeding (nursing) their young, a neocortex (a region of the brain), fur ...
s are imprecise and only colloquial, but historically idealist ideas of transformation and morphology, as in
Goethe Johann Wolfgang von Goethe (28 August 1749 – 22 March 1832) was a German poet, playwright, novelist, scientist, statesman, theatre director, and critic. His works include plays, poetry, literature, and aesthetic criticism, as well as tr ...
's '' Metamorphosis of Plants'', have influenced the development of ideas of
evolution Evolution is change in the heritable characteristics of biological populations over successive generations. These characteristics are the expressions of genes, which are passed on from parent to offspring during reproduction. Variation ...
.


Etymology

The word ''metamorphosis'' derives from Greek , "transformation, transforming", from ('), "after" and ('), "form".


Hormonal control

In insects, growth and metamorphosis are controlled by
hormone A hormone (from the Greek participle , "setting in motion") is a class of signaling molecules in multicellular organisms that are sent to distant organs by complex biological processes to regulate physiology and behavior. Hormones are required ...
s synthesized by
endocrine gland Endocrine glands are ductless glands of the endocrine system that secrete their products, hormones, directly into the blood. The major glands of the endocrine system include the pineal gland, pituitary gland, pancreas, ovaries, testes, thy ...
s near the front of the body ( anterior). Neurosecretory cells in an insect's brain secrete a hormone, the prothoracicotropic hormone (PTTH) that activates prothoracic glands, which secrete a second hormone, usually
ecdysone Ecdysone is a prohormone of the major insect molting hormone 20-hydroxyecdysone, which is secreted from the prothoracic glands. It is of steroidal structure. Insect molting hormones (ecdysone and its homologues) are generally called ecdysteroids. E ...
(an ecdysteroid), that induces
ecdysis Ecdysis is the moulting of the cuticle in many invertebrates of the clade Ecdysozoa. Since the cuticle of these animals typically forms a largely inelastic exoskeleton, it is shed during growth and a new, larger covering is formed. The remnan ...
. PTTH also stimulates the
corpora allata In insect physiology and anatomy, the corpus allatum (plural: corpora allata) is an endocrine gland that generates juvenile hormone; as such, it plays a crucial role in metamorphosis. Surgical removal of the corpora allata (an allatectomy) can cau ...
, a retrocerebral organ, to produce juvenile hormone, which prevents the development of adult characteristics during
ecdysis Ecdysis is the moulting of the cuticle in many invertebrates of the clade Ecdysozoa. Since the cuticle of these animals typically forms a largely inelastic exoskeleton, it is shed during growth and a new, larger covering is formed. The remnan ...
. In holometabolous insects, molts between larval
instar An instar (, from the Latin '' īnstar'', "form", "likeness") is a developmental stage of arthropods, such as insects, between each moult (''ecdysis''), until sexual maturity is reached. Arthropods must shed the exoskeleton in order to grow or ...
s have a high level of juvenile hormone, the moult to the pupal stage has a low level of juvenile hormone, and the final, or imaginal, molt has no juvenile hormone present at all. Experiments on firebugs have shown how juvenile hormone can affect the number of nymph instar stages in hemimetabolous insects. Metamorphosis is iodothyronine-induced and an ancestral feature of all
chordate A chordate () is an animal of the phylum Chordata (). All chordates possess, at some point during their larval or adult stages, five synapomorphies, or primary physical characteristics, that distinguish them from all the other taxa. These fi ...
s.


Insects

All three categories of metamorphosis can be found in the diversity of insects, including no metamorphosis ("ametaboly"), incomplete or partial metamorphosis ("hemimetaboly"), and complete metamorphosis ("holometaboly"). While ametabolous insects show very little difference between larval and adult forms (also known as " direct development"), both hemimetabolous and holometabolous insects have significant morphological and behavioral differences between larval and adult forms, the most significant being the inclusion, in holometabolus organisms, of a
pupa A pupa ( la, pupa, "doll"; plural: ''pupae'') is the life stage of some insects undergoing transformation between immature and mature stages. Insects that go through a pupal stage are holometabolous: they go through four distinct stages in thei ...
l or resting stage between the larval and adult forms.


Development and terminology

In hemimetabolous insects, immature stages are called nymphs. Development proceeds in repeated stages of growth and
ecdysis Ecdysis is the moulting of the cuticle in many invertebrates of the clade Ecdysozoa. Since the cuticle of these animals typically forms a largely inelastic exoskeleton, it is shed during growth and a new, larger covering is formed. The remnan ...
(moulting); these stages are called
instar An instar (, from the Latin '' īnstar'', "form", "likeness") is a developmental stage of arthropods, such as insects, between each moult (''ecdysis''), until sexual maturity is reached. Arthropods must shed the exoskeleton in order to grow or ...
s. The juvenile forms closely resemble adults, but are smaller and lack adult features such as wings and genitalia. The size and morphological differences between nymphs in different instars are small, often just differences in body proportions and the number of segments; in later instars, external wing buds form. The period from one molt to the next is called a stadium. In holometabolous insects, immature stages are called
larva A larva (; plural larvae ) is a distinct juvenile form many animals undergo before metamorphosis into adults. Animals with indirect development such as insects, amphibians, or cnidarians typically have a larval phase of their life cycle. ...
e and differ markedly from adults. Insects which undergo holometabolism pass through a larval stage, then enter an inactive state called
pupa A pupa ( la, pupa, "doll"; plural: ''pupae'') is the life stage of some insects undergoing transformation between immature and mature stages. Insects that go through a pupal stage are holometabolous: they go through four distinct stages in thei ...
(called a "chrysalis" in butterfly species), and finally emerge as adults.


Evolution

The earliest insect forms showed direct development ( ametabolism), and the evolution of metamorphosis in insects is thought to have fuelled their dramatic radiation (1,2). Some early ametabolous "true insects" are still present today, such as bristletails and
silverfish The silverfish (''Lepisma saccharinum'') is a species of small, primitive, wingless insect in the order Zygentoma (formerly Thysanura). Its common name derives from the insect's silvery light grey colour, combined with the fish-like appearance ...
. Hemimetabolous insects include
cockroach Cockroaches (or roaches) are a paraphyletic group of insects belonging to Blattodea, containing all members of the group except termites. About 30 cockroach species out of 4,600 are associated with human habitats. Some species are well-known ...
es, grasshoppers, dragonflies, and
true bugs Hemiptera (; ) is an order of insects, commonly called true bugs, comprising over 80,000 species within groups such as the cicadas, aphids, planthoppers, leafhoppers, assassin bugs, bed bugs, and shield bugs. They range in size from to aroun ...
. Phylogenetically, all insects in the
Pterygota The Pterygota ( grc, πτερυγωτός, pterugōtós, winged) are a subclass of insects that includes the winged insects. It also includes insect orders that are secondarily wingless (that is, insect groups whose ancestors once had wings ...
undergo a marked change in form, texture and physical appearance from immature stage to adult. These insects either have hemimetabolous development, and undergo an incomplete or partial metamorphosis, or holometabolous development, which undergo a complete metamorphosis, including a
pupa A pupa ( la, pupa, "doll"; plural: ''pupae'') is the life stage of some insects undergoing transformation between immature and mature stages. Insects that go through a pupal stage are holometabolous: they go through four distinct stages in thei ...
l or resting stage between the larval and adult forms.Gullan, P.J. & Cranston, P.S. 6.2 ''Life History Patterns and Phases in The Insects: An Outline of Entomology''. pp. 143–153. 2005 by Blackwell Publishing A number of hypotheses have been proposed to explain the evolution of holometaboly from hemimetaboly, mostly centering on whether or not the intermediate stages of hemimetabolous forms are homologous in origin to the pupal stage of holometabolous forms.


Temperature-dependent metamorphosis

According to a 2009 study, temperature plays an important role in insect development as each individual species are found to have specific thermal windows that allow them to progress through their developmental stages. These windows are not significantly affected by ecological traits, rather, the windows are phylogenetically adapted to the ecological circumstances insects are living in.


Recent research

According to research from 2008, adult '' Manduca sexta'' is able to retain behavior learned as a
caterpillar Caterpillars ( ) are the larval stage of members of the order Lepidoptera (the insect order comprising butterflies and moths). As with most common names, the application of the word is arbitrary, since the larvae of sawflies (suborder Sy ...
. Another caterpillar, the ornate moth caterpillar, is able to carry toxins that it acquires from its diet through metamorphosis and into adulthood, where the toxins still serve for protection against predators. Many observations published in 2002, and supported in 2013 indicate that programmed cell death plays a considerable role during physiological processes of multicellular organisms, particularly during
embryogenesis An embryo is an initial stage of development of 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 spe ...
, and metamorphosis. Additional research in 2019 found that both
autophagy Autophagy (or autophagocytosis; from the Ancient Greek , , meaning "self-devouring" and , , meaning "hollow") is the natural, conserved degradation of the cell that removes unnecessary or dysfunctional components through a lysosome-dependent re ...
and
apoptosis Apoptosis (from grc, ἀπόπτωσις, apóptōsis, 'falling off') is a form of programmed cell death that occurs in multicellular organisms. Biochemical events lead to characteristic cell changes ( morphology) and death. These changes in ...
, the two ways programmed cell death occur, are processes undergone during insect metamorphosis. Below is the sequence of steps in the metamorphosis of the butterfly (illustrated):
1 – The larva of a butterfly
2 – The pupa is now spewing the thread to form chrysalis
3 – The chrysalis is fully formed
4 – Adult butterfly coming out of the chrysalis Pieris rapae ''Pieris rapae'' is a small- to medium-sized butterfly species of the whites-and-yellows family Pieridae. It is known in Europe as the small white, in North America as the cabbage white or cabbage butterfly, on several continents as the small c ...
''"> File:ChristianBauer Pieris rapae caterpiller.jpg,
larva A larva (; plural larvae ) is a distinct juvenile form many animals undergo before metamorphosis into adults. Animals with indirect development such as insects, amphibians, or cnidarians typically have a larval phase of their life cycle. ...
File:ChristianBauer Pieris rapae caterpiller2.jpg,
pupa A pupa ( la, pupa, "doll"; plural: ''pupae'') is the life stage of some insects undergoing transformation between immature and mature stages. Insects that go through a pupal stage are holometabolous: they go through four distinct stages in thei ...
File:ChristianBauer Pieris rapae cocoon.jpg, pupa ready to hatch File:ChristianBauer Pieris rapae youngadult.jpg,
adult An adult is a human or other animal that has reached full growth. In human context, the term ''adult'' has meanings associated with social and legal concepts. In contrast to a " minor", a legal adult is a person who has attained the age of maj ...


Chordata


Amphioxus

In cephalochordata, metamorphosis is iodothyronine-induced and it could be an ancestral feature of all
chordate A chordate () is an animal of the phylum Chordata (). All chordates possess, at some point during their larval or adult stages, five synapomorphies, or primary physical characteristics, that distinguish them from all the other taxa. These fi ...
s.


Fish

Some fish, both bony fish (Osteichthyes) and jawless fish (Agnatha), undergo metamorphosis. Fish metamorphosis is typically under strong control by the thyroid hormone. Examples among the non-bony fish include the lamprey. Among the bony fish, mechanisms are varied. The
salmon Salmon () is the common name for several commercially important species of euryhaline ray-finned fish from the family Salmonidae, which are native to tributaries of the North Atlantic (genus '' Salmo'') and North Pacific (genus '' Onco ...
is diadromous, meaning that it changes from a
freshwater Fresh water or freshwater is any naturally occurring liquid or frozen water containing low concentrations of dissolved salts and other total dissolved solids. Although the term specifically excludes seawater and brackish water, it does incl ...
to a saltwater lifestyle. Many species of
flatfish A flatfish is a member of the ray-finned demersal fish order Pleuronectiformes, also called the Heterosomata, sometimes classified as a suborder of Perciformes. In many species, both eyes lie on one side of the head, one or the other migrating ...
begin their life
bilaterally symmetrical Symmetry in biology refers to the symmetry observed in organisms, including plants, animals, fungi, and bacteria. External symmetry can be easily seen by just looking at an organism. For example, take the face of a human being which has a pl ...
, with an eye on either side of the body; but one eye moves to join the other side of the fish – which becomes the upper side – in the adult form. The
European eel The European eel (''Anguilla anguilla'') is a species of eel, a snake-like, catadromous fish. They are normally around and rarely reach more than , but can reach a length of up to in exceptional cases. Eels have been important sources of ...
has a number of metamorphoses, from the larval stage to the
leptocephalus Leptocephalus (meaning "slim head") is the flat and transparent larva of the eel, marine eels, and other members of the superorder Elopomorpha. This is one of the most diverse groups of teleosts, containing 801 species in 4 orders, 24 famili ...
stage, then a quick metamorphosis to glass eel at the edge of the continental shelf (eight days for the Japanese eel), two months at the border of fresh and salt water where the glass eel undergoes a quick metamorphosis into elver, then a long stage of growth followed by a more gradual metamorphosis to the migrating phase. In the pre-adult
freshwater Fresh water or freshwater is any naturally occurring liquid or frozen water containing low concentrations of dissolved salts and other total dissolved solids. Although the term specifically excludes seawater and brackish water, it does incl ...
stage, the eel also has phenotypic plasticity because fish-eating eels develop very wide mandibles, making the head look blunt. Leptocephali are common, occurring in all Elopomorpha ( tarpon- and eel-like fish). Most other bony fish undergo metamorphosis initially from egg to immotile larvae known as ''sac fry'' ( fry with a
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' ...
), then to motile larvae (often known as fingerlings due to them roughly reaching the length of a
human finger A finger is a limb of the body and a type of digit, an organ of manipulation and sensation found in the hands of most of the Tetrapods, so also with humans and other primates. Most land vertebrates have five fingers (Pentadactyly). Chambers ...
) that have to
forage Forage is a plant material (mainly plant leaves and stems) eaten by grazing livestock. Historically, the term ''forage'' has meant only plants eaten by the animals directly as pasture, crop residue, or immature cereal crops, but it is also use ...
for themselves after the yolk sac resorbs, and then to the juvenile stage where the fish progressively start to resemble
adult An adult is a human or other animal that has reached full growth. In human context, the term ''adult'' has meanings associated with social and legal concepts. In contrast to a " minor", a legal adult is a person who has attained the age of maj ...
morphology and behaviors until finally reaching sexual maturity.


Amphibians

In typical amphibian development, eggs are laid in water and larvae are adapted to an aquatic lifestyle. Frogs, toads, and newts all hatch from the eggs as larvae with external gills but it will take some time for the amphibians to interact outside with pulmonary respiration. Afterwards, newt larvae start a predatory lifestyle, while tadpoles mostly scrape food off surfaces with their horny tooth ridges. Metamorphosis in amphibians is regulated by thyroxin concentration in the blood, which stimulates metamorphosis, and
prolactin Prolactin (PRL), also known as lactotropin, is a protein best known for its role in enabling mammals to produce milk. It is influential in over 300 separate processes in various vertebrates, including humans. Prolactin is secreted from the pi ...
, which counteracts its effect. Specific events are dependent on threshold values for different tissues. Because most embryonic development is outside the parental body, development is subject to many adaptations due to specific ecological circumstances. For this reason tadpoles can have horny ridges for teeth, whiskers, and fins. They also make use of the lateral line organ. After metamorphosis, these organs become redundant and will be resorbed by controlled cell death, called
apoptosis Apoptosis (from grc, ἀπόπτωσις, apóptōsis, 'falling off') is a form of programmed cell death that occurs in multicellular organisms. Biochemical events lead to characteristic cell changes ( morphology) and death. These changes in ...
. The amount of adaptation to specific ecological circumstances is remarkable, with many discoveries still being made.


Frogs and toads

With frogs and toads, the external gills of the newly hatched tadpole are covered with a gill sac after a few days, and lungs are quickly formed. Front legs are formed under the gill sac, and hindlegs are visible a few days later. Following that there is usually a longer stage during which the tadpole lives off a vegetarian diet. Tadpoles use a relatively long, spiral‐shaped gut to digest that diet. Recent studies suggest tadpoles don't have a balanced homeostatic feedback control system until the beginning stages of metamorphosis. At this point, their long gut shortens and begins favoring the diet of insects. Rapid changes in the body can then be observed as the lifestyle of the frog changes completely. The spiral‐shaped mouth with horny tooth ridges is resorbed together with the spiral gut. The animal develops a big jaw, and its gills disappear along with its gill sac. Eyes and legs grow quickly, a tongue is formed, and all this is accompanied by associated changes in the neural networks (development of stereoscopic vision, loss of the lateral line system, etc.) All this can happen in about a day, so it is truly a metamorphosis. It is not until a few days later that the tail is reabsorbed, due to the higher thyroxin concentrations required for tail resorption.


Salamanders

Salamander development is highly diverse; some species go through a dramatic reorganization when transitioning from aquatic larvae to terrestrial adults, while others, such as the
axolotl The axolotl (; from nci, āxōlōtl ), ''Ambystoma mexicanum'', is a paedomorphic salamander closely related to the tiger salamander. Axolotls are unusual among amphibians in that they reach adulthood without undergoing metamorphosis. I ...
, display
pedomorphosis Neoteny (), also called juvenilization,Montagu, A. (1989). Growing Young. Bergin & Garvey: CT. is the delaying or slowing of the physiological, or somatic, development of an organism, typically an animal. Neoteny is found in modern humans compared ...
and never develop into terrestrial adults. Within the genus '' Ambystoma'', species have evolved to be pedomorphic several times, and pedomorphosis and complete development can both occur in some species.


Newts

In newts, metamorphosis occurs due to the change in habitat, not a change in diet, because newt larvae already feed as predators and continue doing so as adults. Newts' gills are never covered by a gill sac and will be resorbed only just before the animal leaves the water. Adults can move faster on land than in water. Newts often have an aquatic phase in spring and summer, and a land phase in winter. For adaptation to a water phase,
prolactin Prolactin (PRL), also known as lactotropin, is a protein best known for its role in enabling mammals to produce milk. It is influential in over 300 separate processes in various vertebrates, including humans. Prolactin is secreted from the pi ...
is the required hormone, and for adaptation to the land phase, thyroxin. External gills do not return in subsequent aquatic phases because these are completely absorbed upon leaving the water for the first time.


Caecilians

Basal caecilians such as ''
Ichthyophis ''Ichthyophis'' is a genus of caecilians (limbless amphibians, sometimes called the Asian caecilians) found in Southeast Asia, the southern Philippines The Philippines (; fil, Pilipinas, links=no), officially the Republic of the Philippi ...
'' go through a metamorphosis in which aquatic larva transition into fossorial adults, which involves a loss of the lateral line. More recently diverged caecilians (the Teresomata) do not undergo an
ontogenetic niche shift Ontogenetic niche shift (abbreviated ONS) is an ecological phenomenon where an organism (usually an animal) changes its diet or habitat during its ontogeny (development). During the ontogenetic niche shifting an ecological niche of an individual c ...
of this sort and are in general fossorial throughout their lives. Thus, most caecilians do not undergo an anuran-like metamorphosis.


See also

*
Developmental biology Developmental biology is the study of the process by which animals and plants grow and develop. Developmental biology also encompasses the biology of regeneration, asexual reproduction, metamorphosis, and the growth and differentiation of st ...
* Direct development *
Gosner stage Gosner stage is a generalized system of describing stages of embryonal and larval development in anurans (frogs and toads). The Gosner system includes 46 numbered stages, from fertilized embryo (stage 1) to the completion of metamorphosis (stage ...
* Hypermetamorphosis * Morphogenesis


References


Bibliography

* Davies, R.G. (1998). ''Outlines of Entomology''. Chapman and Hall. Second Edition. Chapter 3. * Williamson D.I. (2003). ''The Origins of Larvae''. Kluwer.


External links

{{Authority control Animal developmental biology Animal anatomy