Metamorphosis

Metamorphosis

Overview
Metamorphosis is a biological process
Biological process
A biological process is a process of a living organism. Biological processes are made up of any number of chemical reactions or other events that results in a transformation....

 by which an animal
Animal
Animals are a major group of multicellular, eukaryotic organisms of the kingdom Animalia or Metazoa. Their body plan eventually becomes fixed as they develop, although some undergo a process of metamorphosis later on in their life. Most animals are motile, meaning they can move spontaneously and...

 physically develops
Developmental biology
Developmental biology is the study of the process by which organisms grow and develop. Modern developmental biology studies the genetic control of cell growth, differentiation and "morphogenesis", which is the process that gives rise to tissues, organs and anatomy.- Related fields of study...

 after birth
Birth
Birth is the act or process of bearing or bringing forth offspring. The offspring is brought forth from the mother. The time of human birth is defined as the time at which the fetus comes out of the mother's womb into the world...

 or hatching, involving a conspicuous and relatively abrupt change in the animal's body
Body
With regard to living things, a body is the physical body of an individual. "Body" often is used in connection with appearance, health issues and death...

 structure through cell growth and differentiation
Cellular differentiation
In developmental biology, cellular differentiation is the process by which a less specialized cell becomes a more specialized cell type. Differentiation occurs numerous times during the development of a multicellular organism as the organism changes from a simple zygote to a complex system of...

. Some insect
Insect
Insects are a class of living creatures within the arthropods that have a chitinous exoskeleton, a three-part body , three pairs of jointed legs, compound eyes, and two antennae...

s, amphibian
Amphibian
Amphibians , are a class of vertebrate animals including animals such as toads, frogs, caecilians, and salamanders. They are characterized as non-amniote ectothermic tetrapods...

s, mollusc
Mollusca
The Mollusca , common name molluscs or mollusksSpelled mollusks in the USA, see reasons given in Rosenberg's ; for the spelling mollusc see the reasons given by , is a large phylum of invertebrate animals. There are around 85,000 recognized extant species of molluscs. Mollusca is the largest...

s, crustacean
Crustacean
Crustaceans form a very large group of arthropods, usually treated as a subphylum, which includes such familiar animals as crabs, lobsters, crayfish, shrimp, krill and barnacles. The 50,000 described species range in size from Stygotantulus stocki at , to the Japanese spider crab with a leg span...

s, Cnidaria
Cnidaria
Cnidaria is a phylum containing over 9,000 species of animals found exclusively in aquatic and mostly marine environments. Their distinguishing feature is cnidocytes, specialized cells that they use mainly for capturing prey. Their bodies consist of mesoglea, a non-living jelly-like substance,...

ns, echinoderm
Echinoderm
Echinoderms are a phylum of marine animals. Echinoderms are found at every ocean depth, from the intertidal zone to the abyssal zone....

s and tunicate
Tunicate
Tunicates, also known as urochordates, are members of the subphylum Tunicata, previously known as Urochordata, a group of underwater saclike filter feeders with incurrent and excurrent siphons that is classified within the phylum Chordata. While most tunicates live on the ocean floor, others such...

s undergo metamorphosis, which is usually accompanied by a change of habitat
Habitat
* Habitat , a place where a species lives and grows*Human habitat, a place where humans live, work or play** Space habitat, a space station intended as a permanent settlement...

 or behavior
Behavior
Behavior or behaviour refers to the actions and mannerisms made by organisms, systems, or artificial entities in conjunction with its environment, which includes the other systems or organisms around as well as the physical environment...

.

Scientific usage of the term is exclusive, and is not applied to general aspects of cell growth
Cell growth
The term cell growth is used in the contexts of cell development and cell division . When used in the context of cell division, it refers to growth of cell populations, where one cell grows and divides to produce two "daughter cells"...

, including rapid growth spurt
Puberty
Puberty is the process of physical changes by which a child's body matures into an adult body capable of reproduction, as initiated by hormonal signals from the brain to the gonads; the ovaries in a girl, the testes in a boy...

s.
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Encyclopedia
Metamorphosis is a biological process
Biological process
A biological process is a process of a living organism. Biological processes are made up of any number of chemical reactions or other events that results in a transformation....

 by which an animal
Animal
Animals are a major group of multicellular, eukaryotic organisms of the kingdom Animalia or Metazoa. Their body plan eventually becomes fixed as they develop, although some undergo a process of metamorphosis later on in their life. Most animals are motile, meaning they can move spontaneously and...

 physically develops
Developmental biology
Developmental biology is the study of the process by which organisms grow and develop. Modern developmental biology studies the genetic control of cell growth, differentiation and "morphogenesis", which is the process that gives rise to tissues, organs and anatomy.- Related fields of study...

 after birth
Birth
Birth is the act or process of bearing or bringing forth offspring. The offspring is brought forth from the mother. The time of human birth is defined as the time at which the fetus comes out of the mother's womb into the world...

 or hatching, involving a conspicuous and relatively abrupt change in the animal's body
Body
With regard to living things, a body is the physical body of an individual. "Body" often is used in connection with appearance, health issues and death...

 structure through cell growth and differentiation
Cellular differentiation
In developmental biology, cellular differentiation is the process by which a less specialized cell becomes a more specialized cell type. Differentiation occurs numerous times during the development of a multicellular organism as the organism changes from a simple zygote to a complex system of...

. Some insect
Insect
Insects are a class of living creatures within the arthropods that have a chitinous exoskeleton, a three-part body , three pairs of jointed legs, compound eyes, and two antennae...

s, amphibian
Amphibian
Amphibians , are a class of vertebrate animals including animals such as toads, frogs, caecilians, and salamanders. They are characterized as non-amniote ectothermic tetrapods...

s, mollusc
Mollusca
The Mollusca , common name molluscs or mollusksSpelled mollusks in the USA, see reasons given in Rosenberg's ; for the spelling mollusc see the reasons given by , is a large phylum of invertebrate animals. There are around 85,000 recognized extant species of molluscs. Mollusca is the largest...

s, crustacean
Crustacean
Crustaceans form a very large group of arthropods, usually treated as a subphylum, which includes such familiar animals as crabs, lobsters, crayfish, shrimp, krill and barnacles. The 50,000 described species range in size from Stygotantulus stocki at , to the Japanese spider crab with a leg span...

s, Cnidaria
Cnidaria
Cnidaria is a phylum containing over 9,000 species of animals found exclusively in aquatic and mostly marine environments. Their distinguishing feature is cnidocytes, specialized cells that they use mainly for capturing prey. Their bodies consist of mesoglea, a non-living jelly-like substance,...

ns, echinoderm
Echinoderm
Echinoderms are a phylum of marine animals. Echinoderms are found at every ocean depth, from the intertidal zone to the abyssal zone....

s and tunicate
Tunicate
Tunicates, also known as urochordates, are members of the subphylum Tunicata, previously known as Urochordata, a group of underwater saclike filter feeders with incurrent and excurrent siphons that is classified within the phylum Chordata. While most tunicates live on the ocean floor, others such...

s undergo metamorphosis, which is usually accompanied by a change of habitat
Habitat
* Habitat , a place where a species lives and grows*Human habitat, a place where humans live, work or play** Space habitat, a space station intended as a permanent settlement...

 or behavior
Behavior
Behavior or behaviour refers to the actions and mannerisms made by organisms, systems, or artificial entities in conjunction with its environment, which includes the other systems or organisms around as well as the physical environment...

.

Scientific usage of the term is exclusive, and is not applied to general aspects of cell growth
Cell growth
The term cell growth is used in the contexts of cell development and cell division . When used in the context of cell division, it refers to growth of cell populations, where one cell grows and divides to produce two "daughter cells"...

, including rapid growth spurt
Puberty
Puberty is the process of physical changes by which a child's body matures into an adult body capable of reproduction, as initiated by hormonal signals from the brain to the gonads; the ovaries in a girl, the testes in a boy...

s. References to "metamorphosis" in mammal
Mammal
Mammals are members of a class of air-breathing vertebrate animals characterised by the possession of endothermy, hair, three middle ear bones, and mammary glands functional in mothers with young...

s are imprecise and only colloquial, but historically idealist ideas of transformation and monadology
Monadology
The Monadology is one of Gottfried Leibniz’s best known works representing his later philosophy. It is a short text which sketches in some 90 paragraphs a metaphysics of simple substances, or monads.- Text :...

, as in Goethe's
Johann Wolfgang von Goethe
Johann Wolfgang von Goethe was a German writer, pictorial artist, biologist, theoretical physicist, and polymath. He is considered the supreme genius of modern German literature. His works span the fields of poetry, drama, prose, philosophy, and science. His Faust has been called the greatest long...

 Metamorphosis of Plants
Metamorphosis of Plants
Johann Wolfgang von Goethe, the great German poet and philosopher published in 1790 the seminal essay Versuch die Metamorphose der Pflanzen zu erklären, known in English as Metamorphosis of Plants. In this work, Goethe essentially discovered the homologous nature of leaf organs in plants, from...

, influenced the development of ideas of evolution
Evolution
Evolution is any change across successive generations in the heritable characteristics of biological populations. Evolutionary processes give rise to diversity at every level of biological organisation, including species, individual organisms and molecules such as DNA and proteins.Life on Earth...

.

Etymology


The word "metamorphosis" derives from Greek
Greek language
Greek is an independent branch of the Indo-European family of languages. Native to the southern Balkans, it has the longest documented history of any Indo-European language, spanning 34 centuries of written records. Its writing system has been the Greek alphabet for the majority of its history;...

 , "transformation, transforming", from (), "change" + (), "form".

Insect metamorphosis


All insects in the Pterygota
Pterygota
Pterygota is a subclass of insects that includes the winged insects. It also includes insect orders that are secondarily wingless ....

 undergo a marked change in form, texture and physical appearance or metamorphosis, from immature 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
Pupa
A pupa is the life stage of some insects undergoing transformation. The pupal stage is found only in holometabolous insects, those that undergo a complete metamorphosis, going through four life stages; embryo, larva, pupa and imago...

l or resting stage between the larval and adult forms.

In hemimetabolous insects, immature stages are called nymphs. Development proceeds in repeated stages of growth and ecdysis
Ecdysis
Ecdysis is the moulting of the cuticula in many invertebrates. This process of moulting is the defining feature of the clade Ecdysozoa, comprising the arthropods, nematodes, velvet worms, horsehair worms, rotifers, tardigrades and Cephalorhyncha...

 (moulting); these stages are called instar
Instar
An instar is a developmental stage of arthropods, such as insects, between each molt , until sexual maturity is reached. Arthropods must shed the exoskeleton in order to grow or assume a new form. Differences between instars can often be seen in altered body proportions, colors, patterns, or...

s. The juvenile forms closely resemble adults, but are smaller and lack adult features such as wings and genitalia. This process is known as "partial" or "incomplete" metamorphosis. The differences between nymphs in different instars are small, often just differences in body proportions and the number of segments, although external wing buds will form in later instars.

In holometabolous insects, immature stages are called larvae, and differ markedly from the adults. Insects which undergo holometabolism pass through a larval stage, then enter an inactive state called pupa
Pupa
A pupa is the life stage of some insects undergoing transformation. The pupal stage is found only in holometabolous insects, those that undergo a complete metamorphosis, going through four life stages; embryo, larva, pupa and imago...

, or chrysalis
Pupa
A pupa is the life stage of some insects undergoing transformation. The pupal stage is found only in holometabolous insects, those that undergo a complete metamorphosis, going through four life stages; embryo, larva, pupa and imago...

, and finally emerge as adults. This process is called "complete" metamorphosis. It is theorized that the pupal stage is the evolutionary compaction of all the nymphal stages of their hemimetabolous ancestors, while the larval stage is an extended, mobile form of the developing embryo.

According to recent research, adult Manduca sexta is able to retain the behaviour learned as a caterpillar
Caterpillar
Caterpillars are the larval form of members of the order Lepidoptera . They are mostly herbivorous in food habit, although some species are insectivorous. Caterpillars are voracious feeders and many of them are considered to be pests in agriculture...

.

Many observations have indicated that programmed cell death
Programmed cell death
Programmed cell-death is death of a cell in any form, mediated by an intracellular program. PCD is carried out in a regulated process which generally confers advantage during an organism's life-cycle...

 plays a considerable role during physiological processes of multicellular organisms, particularly during embryogenesis
Embryogenesis
Embryogenesis is the process by which the embryo is formed and develops, until it develops into a fetus.Embryogenesis starts with the fertilization of the ovum by sperm. The fertilized ovum is referred to as a zygote...

 and metamorphosis.

Hormonal control


Insect
Insect
Insects are a class of living creatures within the arthropods that have a chitinous exoskeleton, a three-part body , three pairs of jointed legs, compound eyes, and two antennae...

 growth and metamorphosis are controlled by hormone
Hormone
A hormone is a chemical released by a cell or a gland in one part of the body that sends out messages that affect cells in other parts of the organism. Only a small amount of hormone is required to alter cell metabolism. In essence, it is a chemical messenger that transports a signal from one...

s synthesized by endocrine glands near the front of the body.

Neurosecretory cells in an insect's brain
Brain
The brain is the center of the nervous system in all vertebrate and most invertebrate animals—only a few primitive invertebrates such as sponges, jellyfish, sea squirts and starfishes do not have one. It is located in the head, usually close to primary sensory apparatus such as vision, hearing,...

 secrete a hormone, the prothoracicotropic hormone
Prothoracicotropic hormone
Prothoracicotropic hormone was the first insect hormone that was discovered. It was originally described simply as "brain hormone" by early workers such as Stefan Kopeć and Vincent Wigglesworth , who realized that ligation of the head of immature insects could prevent molting or pupation of the...

 (PTTH) that activates prothoracic glands, which secrete a second hormone, usually Ecdysone
Ecdysone
Ecdysone is a steroidal prohormone of the major insect molting hormone 20-hydroxyecdysone, which is secreted from the prothoracic glands. Insect molting hormones are generally called ecdysteroids. Ecdysteroids act as moulting hormones of arthropods but also occur in other related phyla where they...

 (a ecdysteroid
Ecdysteroid
Ecdysteroids are insect moulting and sex hormones which include ecdysone and its homologues such as 20-hydroxyecdysone. Ecdysteroids also occur in other invertebrates where they can play a different role. They also appear in many plants mostly as a protection agents against herbivore insects...

), that induces ecdysis.

PTTH also stimulates the corpora allata, a retrocerebral organ, to produce juvenile hormone
Juvenile hormone
Juvenile hormones are a group of acyclic sesquiterpenoids that regulate many aspects of insect physiology. JHs regulate development, reproduction, diapause, and polyphenisms....

 (JH), which prevents the development of adult characteristics during ecdysis
Ecdysis
Ecdysis is the moulting of the cuticula in many invertebrates. This process of moulting is the defining feature of the clade Ecdysozoa, comprising the arthropods, nematodes, velvet worms, horsehair worms, rotifers, tardigrades and Cephalorhyncha...

. In holometabolous insects, molts between larval instars have a high level of JH, the moult to the pupal stage has a low level of JH, and the final, or imaginal, molt has no JH present at all.

Amphibian metamorphosis


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 egg as larvae with external gills. 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
Prolactin also known as luteotropic hormone is a protein that in humans is encoded by the PRL gene.Prolactin is a peptide hormone discovered by Henry Friesen...

, 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
Lateral line
The lateral line is a sense organ in aquatic organisms , used to detect movement and vibration in the surrounding water. Lateral lines are usually visible as faint lines running lengthwise down each side, from the vicinity of the gill covers to the base of the tail...

 organ. After metamorphosis, these organs become redundant and will be resorbed by controlled cell death, called apoptosis
Apoptosis
Apoptosis is the process of programmed cell death that may occur in multicellular organisms. Biochemical events lead to characteristic cell changes and death. These changes include blebbing, cell shrinkage, nuclear fragmentation, chromatin condensation, and chromosomal DNA fragmentation...

. 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.

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 isn't until a few days later that the tail is reabsorbed, due to the higher thyroxin concentrations required for tail resorption.

Newts


In newts, there is no true metamorphosis 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. Just as in tadpoles, their lungs are functional early, but newts don't make as much use of them as tadpoles do. Newts often have an aquatic phase in spring and summer, and a land phase in winter. For adaptation to a water phase, prolactin
Prolactin
Prolactin also known as luteotropic hormone is a protein that in humans is encoded by the PRL gene.Prolactin is a peptide hormone discovered by Henry Friesen...

 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.

Metamorphosis in fish and invertebrate aquatic animals


Little known is that also fish, i.e. both bony fish
Osteichthyes
Osteichthyes , also called bony fish, are a taxonomic group of fish that have bony, as opposed to cartilaginous, skeletons. The vast majority of fish are osteichthyes, which is an extremely diverse and abundant group consisting of over 29,000 species...

 and non-bony fish
Agnatha
Agnatha is a superclass of jawless fish in the phylum Chordata, subphylum Vertebrata. The group excludes all vertebrates with jaws, known as gnathostomes....

 Superclass, undergo metamorphosis. Fish metamorphosis is typically under strong control by thyroid hormone. Examples include the (non-bony fish) Agnatha and lamprey and one bony fish the salmon, which must change from a freshwater to saltwater lifestyle (diadromous). Additionally, the flatfish begins its life bilaterally symmetrical, and one eye must move to join the other side of the fish in its adult form. The European eel
European eel
The European eel, Anguilla anguilla, is a species of eel, a snake-like, catadromous fish. They can reach in exceptional cases a length of 1½ m, but are normally much smaller, about 60–80 cm, and rarely more than 1 m....

 has a number of metamorphoses, from the larval stage to the leptocephalus
Leptocephalus
A leptocephalus is the flat and transparent larva of the eel, marine eels, and other members of the Superorder Elopomorpha. Fishes with a leptocephalus larva stage include the most familiar eels such as the conger, moray eel, and garden eel, and the freshwater eels of the family Anguillidae, plus...

 stage, then a quick outspoken metamorphosis from leptocephalus to glass eel at the edge of the continental shelf (8 days for Japanese eel), two months at the border of fresh and salt water the glass eel undergoes a quick metamorphosis into elver, which then has a long stage of growth followed by a more gradual metamorphosis to the migrating phase. In the pre-adult fresh water stage it also has phenotypic plasticity
Phenotypic plasticity
Phenotypic plasticity is the ability of an organism to change its phenotype in response to changes in the environment. Such plasticity in some cases expresses as several highly morphologically distinct results; in other cases, a continuous norm of reaction describes the functional interrelationship...

 because fish eating eels develop very wide mandibles, making the head look blunt. Leptocephali are very common and a common phase for all Elopomorpha
Elopomorpha
The superorder Elopomorpha contains a variety of types of fishes that range from typical silvery colored fish-like species such as the tarpons and ladyfishes of the Elopiformes and the bonefishes of the Albuliformes, to the long and slender, smooth bodied eels of the Anguilliformes...

 (Tarpon- and eellike fishes).
Most bony fishes undergo metamorphosis after absorption of the yolk sac because after that phase they need to be able to feed for themselves.

See also

  • Hypermetamorphosis
    Hypermetamorphosis
    Hypermetamorphosis is a kind of complete insect metamorphosis in which the different larval instars represent two or more different forms of larva. As the larva molts its morphology can change from that of a campodeiform to scarabaeiform or to vermiform...

  • Morphogenesis
    Morphogenesis
    Morphogenesis , is the biological process that causes an organism to develop its shape...


External links