metamorphosis
Metamorphosis
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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 monadology, as in Goethe's Metamorphosis of Plants, have influenced the development of ideas of evolution.
Contents
Etymology
The word metamorphosis derives from Greek μεταμόρφωσις, "transformation, transforming",[2] from μετα- (meta-), "change" and μορφή (morphe), "form".[3]Hormonal control
Metamorphosis is iodothyronine-induced and an ancestral feature of all chordates.[1]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
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 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 hemimetabolous forms are homologous to pupal form 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.
Other
According to research from 2008, adult Manduca sexta is able to retain behavior learned as a caterpillar.[10] 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.[11]Many observations[when?] have indicated that programmed cell death plays a considerable role during physiological processes of multicellular organisms, particularly during embryogenesis and metamorphosis.
- Sequence illustrating complete metamorphosis in the cabbage white butterfly, ''Pieris rapae''
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pupa ready to hatch
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
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 paedomorphosis and never develop into terrestrial adults. Within the genus Ambystoma, species have evolved to be paedomorphic several times, and paedomorphosis and complete development can both occur in some species.[12]Newts
The large external gills of the crested newt
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.[13] 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.[14]Fish
Some fish, both bony fish (Osteichthyes) and jawless fish (Agnatha), undergo metamorphosis. Fish metamorphosis is typically under strong control by the thyroid hormone.[12]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.[15][16]
See also
References
- Peter B. Moyle and Joseph J. Cech Jr, Fishes: an introduction to ichthyology 5th ed. 9.3: "Development" pp 148ff
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
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. PMID 22174880. doi:10.1371/journal.pone.0028728. 
. PMID 18320055. doi:10.1371/journal.pone.0001736. 
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