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 differentiation.[1] Some insects, jellyfish, fish, amphibians,
mollusks, crustaceans, cnidarians, echinoderms, and tunicates undergo metamorphosis, which is
often accompanied by a change of nutrition source or behavior.[2] Animals can be divided into
species that undergo complete metamorphosis ("holometaboly"), incomplete metamorphosis
("hemimetaboly"), or no metamorphosis ("ametaboly").[3]

Generally organisms with a larval stage undergo metamorphosis, and during metamorphosis the
organism loses larval characteristics.[4]

Etymology

The word metamorphosis derives from Ancient Greek μεταμόρφωσις, "transformation,

transforming",[5] from μετα- (meta-), "after" and μορφή (morphe), "form".[6]

Hormonal control

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 (shedding of the
exoskeleton).[7] 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.[8] Experiments on firebugs have shown how juvenile hormone
can affect the number of nymph instar stages in hemimetabolous insects.[9][10]

In chordates, metamorphosis is iodothyronine-induced and an ancestral feature of all chordates.[11]

Insects

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 holometabolous 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.

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. The period from one molt to the next is called a stadium.[12]

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.[13]

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.[14]

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

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.[15]

Recent research

According to research from 2008, adult Manduca sexta is able to retain behavior learned as a
caterpillar.[16] 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.[17]

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.[18][19] Additional research in 2019 found that both
autophagy and apoptosis, the two ways programmed cell death occur, are processes undergone
during insect metamorphosis. [20]

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

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

 Sequence illustrating complete metamorphosis in the cabbage white butterfly, Pieris rapae

larva

pupa
 

pupa ready for eclosion

adult

Chordata

Amphioxus

In cephalochordata, metamorphosis is iodothyronine-induced and it could be an ancestral feature

of all chordates.[11]

Fish

Some fish, both bony fish (Osteichthyes) and jawless fish (Agnatha), undergo metamorphosis. Fish
metamorphosis is typically under strong control by the thyroid hormone. [21]

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 initially from egg to immotile larvae known as sac
fry (fry with a yolk sac), then to motile larvae (often known as fingerlings due to them roughly
reaching the length of a human finger) that have to forage for themselves after the yolk sac
resorbs, and then to the juvenile stage where the fish progressively start to resemble adult
morphology and behaviors until finally reaching sexual maturity.[22][23]

Amphibians
 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

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. Recent studies suggest tadpoles do not 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.[24]

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. It is not until a few days later that the tail is reabsorbed, due to the higher thyroxin
concentrations required for tail resorption.[25]

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.[21]

Newts

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.[26] 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.[27] 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. [28]