Metamorphosis

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A dragonfly in its final moult, undergoing metamorphosis from its nymph form to an adult Dragonfly metamorphosis.jpg
A dragonfly in its final moult, undergoing metamorphosis from its nymph form to an adult

Metamorphosis is a biological process by which an animal physically develops after birth or hatching, involving a conspicuous and relatively abrupt change in the animal's body structure through cell growth and differentiation. Some insects, fish, amphibians, mollusks, crustaceans, cnidarians, echinoderms, and tunicates undergo metamorphosis, which is often accompanied by a change of nutrition source or behavior. Animals can be divided into species that undergo complete metamorphosis ("holometaboly"), incomplete metamorphosis ("hemimetaboly"), or no metamorphosis ("ametaboly").

Contents

Scientific usage of the term is technically precise, and it is not applied to general aspects of cell growth, including rapid growth spurts. References to "metamorphosis" in mammals are imprecise and only colloquial, but historically idealist ideas of transformation and morphology, as in Goethe's Metamorphosis of Plants , have influenced the development of ideas of evolution.

Etymology

The word metamorphosis derives from Greek μεταμόρφωσις, "transformation, transforming", [1] from μετα- ( meta- ), "after" and μορφή (morphe), "form". [2]

Hormonal control

Metamorphosis is iodothyronine-induced and an ancestral feature of all chordates. [3]

In insects, growth and metamorphosis are controlled by hormones synthesized by endocrine glands 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 (an ecdysteroid), that induces ecdysis. [4] PTTH also stimulates the corpora allata, a retrocerebral organ, to produce juvenile hormone, which prevents the development of adult characteristics during ecdysis. In holometabolous insects, molts between larval instars 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. [5] Experiments on firebugs have shown how juvenile hormone can affect the number of nymph instar stages in hemimetabolous insects. [6] [7]

Insects

Incomplete metamorphosis in the grasshopper with different instar nymphs. The largest specimen is adult. Locust instars NMNS.jpg
Incomplete metamorphosis in the grasshopper with different instar nymphs. The largest specimen is adult.

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 pupal or resting stage between the larval and adult forms.

Development and terminology

Two types of metamorphosis are shown. In a complete (holometabolous) metamorphosis the insect passes through four distinct phases, which produce an adult that does not resemble the larva. In an incomplete (hemimetabolous) metamorphosis an insect does not go through a full transformation, but instead transitions from a nymph to an adult by molting its exoskeleton as it grows. Holometabolous vs. Hemimetabolous.svg
Two types of metamorphosis are shown. In a complete (holometabolous) metamorphosis the insect passes through four distinct phases, which produce an adult that does not resemble the larva. In an incomplete (hemimetabolous) metamorphosis an insect does not go through a full transformation, but instead transitions from a nymph to an adult by molting its exoskeleton as it grows.

In hemimetabolous insects, immature stages are called nymphs. Development proceeds in repeated stages of growth and ecdysis (moulting); these stages are called instars. 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.

In holometabolous insects, immature stages are called larvae and differ markedly from adults. Insects which undergo holometabolism pass through a larval stage, then enter an inactive state called pupa (called a "chrysalis" in butterfly species), and finally emerge as adults. [8]

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. Hemimetabolous insects include cockroaches, grasshoppers, dragonflies, and true bugs. Phylogenetically, all insects in the Pterygota 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 pupal or resting stage between the larval and adult forms. [9]

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.

More recently,[ when? ] scientific attention has turned to characterizing the mechanistic basis of metamorphosis in terms of its hormonal control, by characterizing spatial and temporal patterns of hormone expression relative to metamorphosis in a wide range of insects.

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. [10]

Recent research

According to research from 2008, adult Manduca sexta is able to retain behavior learned as a caterpillar. [11] 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. [12]

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, and metamorphosis. [13] [14]

Below is the sequence of steps in the metamorphosis of the butterfly (illustrated):

Metamorphosis of butterfly (PSF) Metamorphosis of butterfly (PSF).png
Metamorphosis of butterfly (PSF)

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

Chordata

Amphioxus

In cephalochordata, metamorphosis is iodothyronine-induced and it could be an ancestral feature of all chordates. [3]

Fish

Some fish, both bony fish (Osteichthyes) and jawless fish (Agnatha), undergo metamorphosis. Fish metamorphosis is typically under strong control by the thyroid hormone. [15]

Examples among the non-bony fish include the lamprey. Among the bony fish, mechanisms are varied.

The salmon is diadromous, meaning that it changes from a freshwater to a saltwater lifestyle.

Many species of flatfish begin their life bilaterally symmetrical, 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 has a number of metamorphoses, from the larval stage to the leptocephalus 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 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 from embryo to larva (fry) and then to the juvenile stage during absorption of the yolk sac, because after that phase the individual needs to be able to feed for itself. [16] [17]

Amphibians

Just before metamorphosis, only 24 hours are needed to reach the stage in the next picture. RanaTemporariaLarva2.jpg
Just before metamorphosis, only 24 hours are needed to reach the stage in the next picture.
Almost functional common frog with some remains of the gill sac and a not fully developed jaw Rana Temporaria - Larva Final Stage.jpg
Almost functional common frog with some remains of the gill sac and a not fully developed jaw

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, 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. 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 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, display pedomorphosis 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. [15]

Newts

The large external gills of the crested newt LarveKamsalamander.JPG
The large external gills of the crested newt

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. [18] Just as in tadpoles, their lungs are functional early, but newts use them less frequently than tadpoles.[ citation needed ] Newts often have an aquatic phase in spring and summer, and a land phase in winter. For adaptation to a water phase, prolactin 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 go through a metamorphosis in which aquatic larva transition into fossorial adults, which involves a loss of the lateral line. [19] More recently diverged caecilians (the Teresomata) do not undergo an ontogenetic niche shift of this sort and are in general fossorial throughout their lives. Thus, most caecilians do not undergo an anuran-like metamorphosis. [20]

See also

Related Research Articles

Amphibian Class of ectothermic tetrapods

Amphibians are ectothermic, tetrapod 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, arboreal or freshwater aquatic ecosystems. Thus amphibians typically start out as larvae living in water, but some species have developed behavioural adaptations to bypass this.

Neoteny, also called juvenilization, is the delaying or slowing of the physiological development of an organism, typically an animal. Neoteny is found in modern humans. In progenesis, sexual development is accelerated.

Salamander Order of amphibians

Salamanders are a group of amphibians typically characterized by their lizard-like appearance, with slender bodies, blunt snouts, short limbs projecting at right angles to the body, and the presence of a tail in both larvae and adults. All ten extant salamander families are grouped together under the order Urodela. Salamander diversity is highest in the Northern Hemisphere and most species are found in the Holarctic realm, with some species present in the Neotropical realm.

Larva Juvenile form of distinct animals before metamorphosis

A larva 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.

American eel Species of fish

The American eel is a facultative catadromous fish found on the eastern coast of North America. Freshwater eels are fish belonging to the elopomorph superorder, a group of phylogenetically ancient teleosts. The American eel has a slender, snake-like body that is covered with a mucus layer, which makes the eel appear to be naked and slimy despite the presence of minute scales. A long dorsal fin runs from the middle of the back and is continuous with a similar ventral fin. Pelvic fins are absent, and relatively small pectoral fins can be found near the midline, followed by the head and gill covers. Variations exist in coloration, from olive green, brown shading to greenish-yellow and light gray or white on the belly. Eels from clear water are often lighter than those from dark, tannic acid streams.

Pupa Life stage of some insects undergoing transformation

A pupa 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 their life cycle, the stages thereof being egg, larva, pupa, and imago. The processes of entering and completing the pupal stage are controlled by the insect's hormones, especially juvenile hormone, prothoracicotropic hormone, and ecdysone. The act of becoming a pupa is called pupation, and the act of emerging from the pupal case is called eclosion or emergence.

Aquatic animal Animal that lives in water for most or all of its lifetime

An aquatic animal is any animal, whether invertebrate or vertebrate, that lives in water for most or all of its lifetime. Many insects such as mosquitoes, mayflies, dragonflies and caddisflies have aquatic larvae, with winged adults. Aquatic animals may breathe air or extract oxygen from water through specialised organs called gills, or directly through the skin. Natural environments and the animals that live in them can be categorized as aquatic (water) or terrestrial (land). This designation is polyphyletic.

Instar Developmental stage of arthropods between moults

An instar 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 assume a new form. Differences between instars can often be seen in altered body proportions, colors, patterns, changes in the number of body segments or head width. After moulting, i.e. shedding their exoskeleton, the juvenile arthropods continue in their life cycle until they either pupate or moult again. The instar period of growth is fixed; however, in some insects, like the salvinia stem-borer moth, the number of instars depends on early larval nutrition. Some arthropods can continue to moult after sexual maturity, but the stages between these subsequent moults are generally not called instars.

Northern crested newt Species of amphibian

The northern crested newt, great crested newt or warty newt is a newt species native to Great Britain, northern and central continental Europe and parts of Western Siberia. It is a large newt, with females growing up to 16 cm (6.3 in) long. Its back and sides are dark brown, while the belly is yellow to orange with dark blotches. Males develop a conspicuous jagged crest on their back and tail during the breeding season.

In animal dormancy, diapause is the delay in development in response to regularly and recurring periods of adverse environmental conditions. It is a physiological state with very specific initiating and inhibiting conditions. The mechanism is a means of surviving predictable, unfavorable environmental conditions, such as temperature extremes, drought, or reduced food availability. Diapause is observed in all the life stages of arthropods, especially insects. Embryonic diapause, a somewhat similar phenomenon, occurs in over 130 species of mammals, possibly even in humans, and in the embryos of many of the oviparous species of fish in the order Cyprinodontiformes.

Veliger

A veliger is the planktonic larva of many kinds of sea snails and freshwater snails, as well as most bivalve molluscs (clams) and tusk shells.

Holometabolism, also called complete metamorphosis, is a form of insect development which includes four life stages: egg, larva, pupa, and imago. Holometabolism is a synapomorphic trait of all insects in the superorder Endopterygota. Immature stages of holometabolous insects are very different from the mature stage. In some species the holometabolous life cycle prevents larvae from competing with adults because they inhabit different ecological niches. The morphology and behavior of each stage are adapted for different activities. For example, larval traits maximize feeding, growth, and development, while adult traits enable dispersal, mating, and egg laying. Some species of holometabolous insects protect and feed their offspring. Other insect developmental strategies include ametabolism and hemimetabolism.

Polyphenism Type of polymorphism where different forms of an animal arise from a single genotype

A polyphenic trait is a trait for which multiple, discrete phenotypes can arise from a single genotype as a result of differing environmental conditions. It is therefore a special case of phenotypic plasticity.

Juvenile (organism) Individual organism that has not yet reached its adult form

A juvenile is an individual organism that has not yet reached its adult form, sexual maturity or size. Juveniles can look very different from the adult form, particularly in colour, and may not fill the same niche as the adult form. In many organisms the juvenile has a different name from the adult.

Imaginal disc One of the parts of a holometabolous insect larva

An imaginal disc is one of the parts of a holometabolous insect larva that will become a portion of the outside of the adult insect during the pupal transformation. Contained within the body of the larva, there are pairs of discs that will form, for instance, the wings or legs or antennae or other structures in the adult. The role of the imaginal disc in insect development was first elucidated by Jan Swammerdam.

<i>Cotesia congregata</i> Species of wasp

Cotesia congregata is a parasitoid wasp of the genus Cotesia. The genus is particularly noted for its use of polydnaviruses. Parasitoids are distinct from true parasites in that a parasitoid will ultimately kill its host or otherwise sterilize it.

Juvenile-hormone esterase

In enzymology, juvenile hormone esterase (JH esterase) is an enzyme that catalyzes the hydrolysis of juvenile hormone. For example, the juvenile hormone II (found in Lepidoptera):

Newt Salamander in the subfamily Pleurodelinae

A newt is a salamander in the subfamily Pleurodelinae. The terrestrial juvenile phase is called an eft. Unlike other members of the family Salamandridae, newts are semiaquatic, alternating between aquatic and terrestrial habitats. Not all aquatic salamanders are considered newts, however. More than 100 known species of newts are found in North America, Europe, North Africa and Asia. Newts metamorphose through three distinct developmental life stages: aquatic larva, terrestrial juvenile (eft), and adult. Adult newts have lizard-like bodies and return to the water every year to breed, otherwise living in humid, cover-rich land habitats.

Neurosecretion is the storage, synthesis and release of hormones from neurons. These neurohormones, produced by neurosecretory cells, are normally secreted from nerve cells in the brain that then circulate into the blood. These neurohormones are similar to nonneural endocrine cells and glands in that they also regulate both endocrine and nonendocrine cells. Neurosecretion cells synthesize and package their product in vesicles and exocytose them at axon endings just as normal neurons do, but release their product farther from their target than normal neurons, typically releasing their neurohormones into the circulatory system to reach their distant targets.

Juvenile fish

Juvenile fish go through various stages between birth and adulthood. They start as eggs which hatch into larvae. The larvae are not able to feed themselves, and carry a yolk-sac which provides their nutrition. Before the yolk-sac completely disappears, the tiny fish must become capable of feeding themselves. When they have developed to the point where they are capable of feeding themselves, the fish are called fry. When, in addition, they have developed scales and working fins, the transition to a juvenile fish is complete and it is called a fingerling. Fingerlings are typically about the size of fingers. The juvenile stage lasts until the fish is fully grown, sexually mature and interacting with other adult fish.

References

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Bibliography

Commons-logo.svg Media related to metamorphosis at Wikimedia Commons

Wiktionary-logo-en-v2.svg The dictionary definition of metamorphosis at Wiktionary