![Cicada molting animated-2](https://upload.wikimedia.org/wikipedia/commons/6/62/Cicada_molting_animated-2.gif)
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
An exoskeleton (from Greek ''éxō'' "outer" and ''skeletós'' "skeleton") is an external skeleton that supports and protects an animal's body, in contrast to an internal skeleton (endoskeleton) in for example, a human. In usage, some of the ...
, it is shed during growth and a new, larger covering is formed.
The remnants of the old, empty exoskeleton are called
exuviae
In biology, exuviae are the remains of an exoskeleton and related structures that are left after ecdysozoans (including insects, crustaceans and arachnids) have moulted. The exuviae of an animal can be important to biologists as they can often b ...
.
After moulting, an arthropod is described as ''teneral'', a ''callow''; it is "fresh", pale and soft-bodied. Within one or two hours, the cuticle hardens and darkens following a
tanning
Tanning may refer to:
*Tanning (leather), treating animal skins to produce leather
*Sun tanning, using the sun to darken pale skin
**Indoor tanning, the use of artificial light in place of the sun
**Sunless tanning, application of a stain or dye t ...
process analogous to the production of
leather
Leather is a strong, flexible and durable material obtained from the tanning, or chemical treatment, of animal skins and hides to prevent decay. The most common leathers come from cattle, sheep, goats, equine animals, buffalo, pigs and hogs, ...
.
During this short phase the animal expands, since growth is otherwise constrained by the rigidity of the exoskeleton. Growth of the limbs and other parts normally covered by the hard exoskeleton is achieved by transfer of body fluids from soft parts before the new skin hardens. A spider with a small abdomen may be undernourished but more probably has recently undergone ecdysis. Some arthropods, especially large insects with tracheal respiration, expand their new exoskeleton by swallowing or otherwise taking in air. The maturation of the structure and
colouration of the new exoskeleton might take days or weeks in a long-lived insect; this can make it difficult to identify an individual if it has recently undergone ecdysis.
Ecdysis allows damaged tissue and missing limbs to be
regenerated or substantially re-formed. Complete regeneration may require a series of moults, the stump becoming a little larger with each moult until it is a normal, or near normal, size.
Etymology
The term ''ecdysis'' comes from
Ancient Greek
Ancient Greek includes the forms of the Greek language used in ancient Greece and the ancient world from around 1500 BC to 300 BC. It is often roughly divided into the following periods: Mycenaean Greek (), Dark Ages (), the Archaic peri ...
() 'to take off, strip off'.
Process
In preparation for ecdysis, the arthropod becomes inactive for a period of time, undergoing
apolysis
Apolysis ( grc, ἀπόλυσις "discharge, lit. absolution") is the separation of the cuticle from the epidermis in arthropods and related groups (Ecdysozoa). Since the cuticle of these animals is also the skeletal support of the body and is in ...
or separation of the old exoskeleton from the underlying epidermal cells. For most organisms, the resting period is a stage of preparation during which the secretion of fluid from the moulting glands of the epidermal layer and the loosening of the underpart of the cuticle occurs.
Once the old cuticle has separated from the epidermis, a digesting fluid is secreted into the space between them. However, this fluid remains inactive until the upper part of the new cuticle has been formed. Then, by crawling movements, the organism pushes forward in the old
integumentary shell, which splits down the back allowing the animal to emerge. Often, this initial crack is caused by a combination of movement and increase in blood pressure within the body, forcing an expansion across its
exoskeleton
An exoskeleton (from Greek ''éxō'' "outer" and ''skeletós'' "skeleton") is an external skeleton that supports and protects an animal's body, in contrast to an internal skeleton (endoskeleton) in for example, a human. In usage, some of the ...
, leading to an eventual crack that allows for certain organisms such as
spider
Spiders ( order Araneae) are air-breathing arthropods that have eight legs, chelicerae with fangs generally able to inject venom, and spinnerets that extrude silk. They are the largest order of arachnids and rank seventh in total species ...
s to extricate themselves.
While the old cuticle is being digested, the new layer is secreted. All cuticular structures are shed at ecdysis, including the inner parts of the exoskeleton, which includes terminal linings of the
alimentary tract
The gastrointestinal tract (GI tract, digestive tract, alimentary canal) is the tract or passageway of the digestive system that leads from the mouth to the anus. The GI tract contains all the major organs of the digestive system, in humans and ...
and of the
trachea
The trachea, also known as the windpipe, is a Cartilage, cartilaginous tube that connects the larynx to the bronchi of the lungs, allowing the passage of air, and so is present in almost all air-breathing animals with lungs. The trachea extends ...
e if they are present.
Insects
Each stage of development between moults for insects in the taxon
endopterygota is called an
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 ...
, or stadium, and each stage between moults of insects in the
Exopterygota
The Exopterygota (Ancient Greek ''ἔξω'' (éxō, “outside”) + ''πτερόν'' (pterón, “wing”) + New Latin ''-ota'' (“having”)), also known as Hemipterodea, are a superorder of insects of the subclass Pterygota in the infracla ...
is called a nymph: there may be up to 15 nymphal stages.
Endopterygota tend to have only four or five instars. Endopterygotes have more alternatives to moulting, such as expansion of the cuticle and collapse of air sacs to allow growth of internal organs.
The process of moulting in insects begins with the separation of the cuticle from the underlying epidermal cells (apolysis) and ends with the shedding of the old cuticle (ecdysis). In many species it is initiated by an increase in the
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 ...
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. ...
. This hormone causes:
* apolysis – the separation of the cuticle from the
epidermis
*
secretion of new cuticle materials beneath the old
* degradation of the old cuticle
After apolysis the insect is known as a
pharate
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 their ...
. Moulting fluid is then secreted into the exuvial space between the old cuticle and the epidermis, this contains inactive enzymes which are activated only after the new
epicuticle
The cuticle forms the major part of the integument of the Arthropoda. It includes most of the material of the exoskeleton of the insects, Crustacea, Arachnida, and Myriapoda.
Morphology
In arthropods, the integument, the external "skin", or ...
is secreted. This prevents the new
procuticle
The cuticle forms the major part of the integument of the Arthropoda. It includes most of the material of the exoskeleton of the insects, Crustacea, Arachnida, and Myriapoda.
Morphology
In arthropods, the integument, the external "skin", or "s ...
from getting digested as it is laid down. The lower regions of the old cuticle, the
endocuticle
The cuticle forms the major part of the integument of the Arthropoda. It includes most of the material of the exoskeleton of the insects, Crustacea, Arachnida, and Myriapoda.
Morphology
In arthropods, the integument, the external "skin", or ...
and
mesocuticle
The cuticle forms the major part of the integument of the Arthropoda. It includes most of the material of the exoskeleton of the insects, Crustacea, Arachnida, and Myriapoda.
Morphology
In arthropods, the integument, the external "skin", or ...
, are then digested by the enzymes and subsequently absorbed. The
exocuticle
The cuticle forms the major part of the integument of the Arthropoda. It includes most of the material of the exoskeleton of the insects, Crustacea, Arachnida, and Myriapoda.
Morphology
In arthropods, the integument, the external "skin", or ...
and epicuticle resist digestion and are hence shed at ecdysis.
Spiders
Spiders generally change their skin for the first time while still inside the egg sac, and the spiderling that emerges broadly resembles the adult. The number of moults varies, both between species and sexes, but generally will be between five times and nine times before the spider reaches maturity. Not surprisingly, since males are generally smaller than females, the males of many species mature faster and do not undergo ecdysis as many times as the females before maturing.
Members of the
Mygalomorphae
The Mygalomorphae, or mygalomorphs, are an infraorder of spiders, and comprise one of three major groups of living spiders with over 3000 species, found on all continents except Antarctica. Many members are known as trapdoor spiders due to the ...
are very long-lived, sometimes 20 years or more; they moult annually even after they mature.
Spiders stop feeding at some time before moulting, usually for several days. The physiological processes of releasing the old exoskeleton from the tissues beneath typically cause various colour changes, such as darkening. If the old exoskeleton is not too thick it may be possible to see new structures, such as
seta
In biology, setae (singular seta ; from the Latin word for " bristle") are any of a number of different bristle- or hair-like structures on living organisms.
Animal setae
Protostomes
Annelid setae are stiff bristles present on the body. ...
e, from the outside. However, contact between the nerves and the old exoskeleton is maintained until a very late stage in the process.
The new, teneral exoskeleton has to accommodate a larger frame than the previous
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 ...
, while the spider has had to fit into the previous exoskeleton until it has been shed. This means the spider does not fill out the new exoskeleton completely, so it commonly appears somewhat wrinkled.
Most species of spiders hang from silk during the entire process, either dangling from a drop line, or fastening their claws into webbed fibres attached to a suitable base. The discarded, dried exoskeleton typically remains hanging where it was abandoned once the spider has left.
To open the old exoskeleton, the spider generally contracts its abdomen (
opisthosoma
The opisthosoma is the posterior part of the body in some arthropods, behind the prosoma (cephalothorax). It is a distinctive feature of the subphylum Chelicerata (arachnids, horseshoe crabs and others). Although it is similar in most respects to a ...
) to supply enough fluid to pump into the
prosoma
The cephalothorax, also called prosoma in some groups, is a tagma of various arthropods, comprising the head and the thorax fused together, as distinct from the abdomen behind. (The terms ''prosoma'' and ''opisthosoma'' are equivalent to ''cepha ...
with sufficient pressure to crack it open along its lines of weakness. The
carapace
A carapace is a Dorsum (biology), dorsal (upper) section of the exoskeleton or shell in a number of animal groups, including arthropods, such as crustaceans and arachnids, as well as vertebrates, such as turtles and tortoises. In turtles and tor ...
lifts off from the front, like a helmet, as its surrounding skin ruptures, but it remains attached at the back. Now the spider works its limbs free and typically winds up dangling by a new thread of silk attached to its own exuviae, which in turn hang from the original silk attachment.
At this point the spider is a callow; it is teneral and vulnerable. As it dangles, its exoskeleton hardens and takes shape. The process may take minutes in small spiders, or some hours in the larger Mygalomorphs. Some spiders, such as some ''
Synema'' species, members of the
Thomisidae
The Thomisidae are a family of spiders, including about 170 genera and over 2,100 species. The common name crab spider is often linked to species in this family, but is also applied loosely to many other families of spiders. Many members of th ...
(crab spiders), mate while the female is still callow, during which time she is unable to eat the male.
Eurypterids
Eurypterid
Eurypterids, often informally called sea scorpions, are a group of extinct arthropods that form the order Eurypterida. The earliest known eurypterids date to the Darriwilian stage of the Ordovician period 467.3 million years ago. The group is l ...
s are a group of
chelicerates
The subphylum Chelicerata (from New Latin, , ) constitutes one of the major subdivisions of the phylum Arthropoda. It contains the sea spiders, horseshoe crabs, and arachnids (including harvestmen, scorpions, spiders, solifuges, ticks, and mit ...
that became
extinct
Extinction is the termination of a kind of organism or of a group of kinds (taxon), usually a species. The moment of extinction is generally considered to be the death of the last individual of the species, although the capacity to breed and ...
in the
Late Permian
Late may refer to:
* LATE, an acronym which could stand for:
** Limbic-predominant age-related TDP-43 encephalopathy, a proposed form of dementia
** Local-authority trading enterprise, a New Zealand business law
** Local average treatment effect, ...
. They underwent ecdysis similarly to extant chelicerates, and most fossils are thought to be of exuviae, rather than cadavers.
See also
*
Ecdysteroid
Ecdysteroids are arthropod steroid hormones that are mainly responsible for molting, development and, to a lesser extent, reproduction; examples of ecdysteroids include ecdysone, ecdysterone, turkesterone and 2-deoxyecdysone. These compounds are ...
References
External links
*{{Commons category-inline
Animal developmental biology
Protostome anatomy
Ethology
it:Muta (biologia)