The term neoteny is derived from Latin neotenia; neo + Greek teinien = to

extend, meaning when ​larval life is extended.​ Neoteny also called

juvenilization, is the ​retention, by adults in a species, of traits previously
seen only in juveniles.​ Neoteny has two forms depending upon the capability of
individuals to breed.

Pedomorphosis was first proposed by Walter Garstang in 1922. The underlying

mechanisms for this include heterochrony (change in features during
development). It is common in many animal species domesticated by humans,
including dogs, chickens, pigs and cattle. It is believed to be a side-effect of the
selective pressure of human-directed breeding ​for juvenile behavioral
characteristics such as docility and cuteness. Pedomorphosis also occurs in
termites and several species of cockroach. Humans are considered by some
scientists to be pedomorphic, due to their flattened face, short jaw, and bulbous
forehead compared to other adult primates.

Paedogenesis is the act of reproduction by an organism that has not achieved

physical maturity. In other words paedogenesis is the production of offspring by
an organism in its larval or juvenile form and elimination of the adult phase of the
life cycle. It is associated with progenesis, where sexual maturity is achieved in
the juvenile form and further physical maturity is not achieved. Paedogenesis is
found in insects in which the larval stage reproduces without achieving maturity.
It occurs in the females of certain beetles, Strepsiptera, bagworms, scale insects
and gall midges.

EXAMPLES OF NEOTENY

Flightlessbirds
Flightless birds such as ostriches, emus, cassowaries and kiwis are believed to
have evolved by retaining characteers of chicks and losing ability to fly. Physical
proportions of these birds resemble those of the chicks of flighted birds.

Humans

With human traits such as sparse body hairs and enlarged heads are thought to
be reminiscent of baby primates. Lactose tolerance in adults is a form of neoteny
now considered normal in certain populations that traditionally consume milk
while most other humans are lactose intolerant as adults. Although females
mature at an earlier age, women do not go on to acquire the toughened skin,
coarse body hair, thyroid cartilage, bony eye ridges, or deepened voices which
are the common inheritance of most adult hominoids and other primates.
Paedomorphic characteristics in women are widely acknowledged as desirable
by men. For instance, vellus hair is a juvenile characteristic. However, while men
develop longer, coarser, thicker, and darker terminal hair through sexual
differentiation, women do not, leaving their vellus hair visible.

Stephen Jay Gould was an advocate of the view that humans are a neotenous
species of chimpanzee. The argument is that juvenile chimpanzees have an
almost-identical bone structure to humans, and that the chimpanzee’s ability to
learn seems to be cut off upon reaching maturity.

Another theory suggests that humans’ neotenous characteristics were an

evolutionary strategy that enabled Cro-Magnons to gain predominance over
neanderthals (and possibly H. erectus and H. heidelbergensis) by appealing to
these species’ nurturing instincts through paedomorphic cuteness and to avoid
territorial aggression.

Bolk (1926) provided an abbreviated list of human neotenic characters:

Our “flat faced” orthognathy.

Reduction or lack of body hair.
Loss of pigmentation in skin, eyes, and hair.
The form of the external ear.
The central position of the foramen magnum (it migrates backward during the
ontogeny of primates).
High relative brain weight.
Persistence of the cranial sutures to an advanced age.
The labia majora of women.
The structure of the hand and foot.
The form of the pelvis.
The ventrally directed position of the sexual canal in women.
Small teeth and variations of the tooth row and cranial sutures. Late eruption of
teeth.
Absence of eye brow ridges.
Absence of cranial crests.
Thinness of skull bones.
Position of orbits under cranial cavity.
No rotation of the big toe.
Prolonged period of infantile dependency and prolonged period of growth.
Long life span.
Crying is among the child-like behaviors. Certainly human facial anatomy and
physiology are intricately engineered for both the discharge of tears and the
facial and vocal expressions that accompany them to draw attention and
sympathy.
Origin of chordates

It is possible that chordates originated by neoteny. Molecular evidence suggests

that the nearest relatives of the chordates are the tunicates, marine filter feeders.
Although sessile in their adult, sexually mature form, tunicates have a motile
larval dispersal form, which has a notochord similar to that found in chordates. At
some point, the motile larvae of the tunicate became sexually mature before
metamorphosis and gave rise to free swimming chordates.

Neoteny in amphibia

Natural pedomorphosis occurs in many species of amphibians, especially

ambystomatid and protean salamanders. In amphibians it can be obligate or
facultative. For example, some salamanders retain their gills during adulthood,
unlike most other amphibians.

Most neotenic populations belong to the Tiger Salamander complex –

Ambystoma tigrinum, Ambystoma velasci, Ambystoma mavortium, and their
close relatives. Wholly neotenic Ambystoma species include the Axolotl,
Ambystoma taylori, Ambystoma andersoni, and Ambystoma dumerilii. Neotenes
retain ability to regenerate limbs, tails, and nearly every organ in their body.
Neoteny, sometimes called paedomorphism, is apparent in Urodela, except
Rhyacotritonidae.

There are three types of neoteny, obligate, inducible obligate, and facultative.

Obligate neoteny

All members of the families Amphiumidae, Sirenidae, Cryptobranchidae, and

Proteidae are obligate neotenes, meaning they never fully metamorphose, and
retain larval characteristics in varying degrees into adulthood. Most of these
species are insensitive to thyroid hormone doses.

Inducible obligate neoteny

Inducible obligate neotenes of the family Ambystomatidae, and some species of

the family Plethodontidae are unique in that metamorphosis can be induced by
manipulating the thyroid function in the laboratory. The most famous inducible
obligate neotene is perhaps Ambystoma mexicanum (Ambystomatidae).

Facultative neoteny

Facultative neoteny is observed in Salamandridae, Dicamptodontidae,

Ambystomatidae, Hynobiidae, and Plethodontidae. Facultative neoteny occurs in
individuals or entire populations as a result of environmental factors. For
example, in extremely cold temperatures, where a terrestrial existence would be
inhospitable, individuals or populations may remain aquatic, retaining larval
characteristics into adulthood.

The Axolotl larva and Neoteny

A sexually mature axolotl, at age 18–24 months, ranges in length from 15–45
centimetres. Axolotls possess features typical of salamander larvae, including
external gills and a caudal fin extending from behind the head to the vent. Their
heads are wide, and their eyes are lidless. Their limbs are underdeveloped and
possess long, thin digits. Males are identified by their swollen cloaca lined with
papillae, while females are noticeable for their wider bodies full of eggs. Axolotls
have barely visible vestigial teeth which would have developed during
metamorphosis. Three pairs of external gills are used for respiration, although
buccal pumping (gulping air from the surface) may also be used in order to
provide oxygen to their lungs.

2. Types of Neoteny:
Kollmann (1882) classified neoteny into two categories, viz., Partial neoteny and
Total neoteny.

i. Partial neoteny:

Partial neoteny involves the simple postponement of metamorphosis beyond the

normal period due to temporary change in ecological condition or due to sudden
physiological abnormality.

Wintering of the tadpdles of Pelobates fuscus, Pelodytes punctatus, Alytes

obstetricans, Hyla arborea, Bufo vulgaris, Bufo viridis, Rana temporaria, Rana
esculenta, Bombinator pachypus and many others furnish the typical examples of
partial neoteny.

In Alytes the broods usually complete their development within autumn. But the
larvae which hatched later in the months of July or August usually hibernate and
retain their larval features up to the next autumn.
In Rana esculenta most of the larvae attain much larger size than the normal and
remain in an under-developed stage, i.e., the hind limbs reach the budding stage,
the head and the body attains a blotted and unhealthy look. Majority of the larvae
remain in this slug​gish stage for one or two years after which metamorphosis
takes place.

ii. Total neoteny

In this category the speci​mens become sexually mature at the larval stage but
retain larval characters, like (i) exter​nal gills, (ii) tail-fin, (iii) ill-developed eyes, (iv)
ill-developed fin on the back and (v) very weak limbs. Total neotenic animals are
paedo-genic.

Because paedogenesis involves the capability of reproduction at the larval stage.

In this case of total neoteny the sexually functional larvae cease to
metamorphosis. In extreme cases of total neoteny (e.g., Necturus, Proteus, etc.)
the larvae attain sexual maturity and they remain in that stage without
undergoing metamorphosis.

Intermediate stages between partial and total neoteny are also recorded where
the lar​vae become sexually functional and may metamorphose into adults with
the advent of favourable conditions. The sexually mature axolotl larvae come
under this category.

Total neoteny is seen in many urodele (e.g., Necturus, Amphiuma, Triturus

vulgaris, Triton cristatus, Triton waltli, Ambystoma, Triturus alpestris, Siren,
Proteus, etc.). The phe​nomenon of neoteny has been extensively studied in case
of Ambystoma which, however, does not show an extreme case of neoteny. The
axolotl larvae of Ambystoma can metamor​phose into adult Ambystoma under
favourable conditions.

Factors affecting neoteny

In the axolotl, metamorphic failure is caused by a lack of thyroid stimulating

hormone, which is used to induce the thyroid to produce ​thyroxine​ in
transforming salamanders. Axolotls can be induced to metamorphose by an
injection of iodine​ (used in the production of thyroid hormones) or by shots of
thyroxine hormone.

Another method for inducing metamorphosis is to keep them in ​shallow water

tanks.​ They will then, over a period of weeks, slowly metamorphose into adult
salamanders. However, most attempts at inducing metamorphosis lead to death.
This is likely due to the strong genetic basis for neoteny in pet axolotls. Artificial
metamorphosis also dramatically shortens the axolotl’s lifespan, if they survive
the process.

Research indicates that neoteny occurs because the hypothalamus of the brain
fails to produce the hormone that causes the pituitary to stimulate the thyroid
gland to produce growth hormones that trigger metamorphosis. Some scientists
think that neoteny may have evolved as a response to the hazards of life on land.

Extrinsic factors:
● Gadow (1901) advanced the idea that the cause of retention of larval
features in axolotl is the abundance of food and other favourable requisites
in aquatic life.
● Shufeldt holds that deep water and cold​ness inhibit thyroxine secretion
which retards metamorphosis.
● Drying up of swamps, lack of adequate food supply and rise in temperature
in surrounding water induce metamorphosis.
● Weissmann again claimed that the retardation of metamorphosis of the
axolotls is possibly due to the saline nature of the water of the lakes where
they live.
● In an investigation to establish the role of physical factors in neoteny,
tadpoles were kept in water-holes with high vertical walls, so that they were
not allowed to reach the land above the water-holes. It has been observed
that this forced and prolonged use of larval gills and tails cause their
further development, whereas the growth of limbs and other structures
nec​essary for terrestrial life remained suspended.
● It has further been observed that the axolotls which were not likely to
metamor​phose in normal habitat could be forced to metamorphose by
slowly accustoming them to land-life.
 ● Reverse phenomenon also happened in the life of axolotls. Partly
metamorphosed axolotls accustomed to land-life could again induced to
return to larval life.
● Huxley (1929) undertook an investigation on temperature-coefficient on
metamorphosis. Several half-grown anuran tadpoles were cul​tured in
variable temperatures ranging from 3-30°C for same span of time. But the
culture solution has same concentration of thyroxine.
● The larvae exposed to temperature range below 5°C could not complete
their metamor​phosis even when exposed to higher tempe​rature. But the
larvae exposed to higher temperature from the beginning completed
metamorphosis very quickly.

Intrinsic factors influencing neoteny:

● Zondeck and Leiter (1923) established that ​calcium delays
metamorphosis​ in axolotls. Gressner (1928) also advanced that insulin
hormone inhibits metamorphosis.
● But recent researches incline to reveal that the metamorphosis is primarily
influenced (i) by varying threshold levels of thyroxine and its analogs and
(ii) by the degree of responsiveness of the larval tissue to hormones.
● During early pre-metamorphic stage in amphibian development, the level
of thyroxine (T4) is kept very low in the body by genetic mechanism. Etkin
and his colla​borators have also established the role of pro​lactin on
metamorphosis.
● They have shown that the level of prolactin which acts as an inhibitor in the
overall control of metamorpho​sis remains high at this time. In the light of
modern genetics it may be suggested that the structural genes guiding the
synthesis of thy​roxine are ‘switched off’ by some operator genes whereas
the genes guiding the formation of prolactin are ‘switched on’.
● In such condi​tion the hypothalamus becomes sensitive to the available
level of thyroid hormone in the blood stream. The neurosecretory
apparatus of the hypothalamus produces a substance, called
thyrotropin-releasing factor (TRF). TRF stimu​lates the anterior lobe of the
pituitary to produce thyroid-stimulating hormone (TSH) which in turn
enhances the rate of thyroid secretion.
● As the level of TSH rises during pro-metamorphosis, the level of prolactin
suddenly falls. So the metamorphosis starts. The time of shift in hor​mone
 balance is possibly determined by the initiation of positive thyroid
feed-back to the hypothalamus. Poor secretion of thyroid glands and the
irresponsiveness of the larval tissues to the hormone are responsible for
neoteny.
● Kuhn (1925) studied the thyroid glands of neotenous larvae of the warty
newt and observed that the alveoli which secrete thy​roxine remain
undeveloped and even axolotls possessing normally developed thyroid
glands failed to pour their secretion in the blood stream.
● Transplantation of at least two more thyroid glands in addition to the
nor​mal glands causes the metamorphosis of axolotls. This result indicated
that the thyroid gland of axolotls is able to produce one-third of the
required quantity of thyroxine and that the larval tissues are only one-third
responsive in contrast to normal specimens.
● Bytenski and Saez have experimentally exchanged the pituitary gland
between a sala​mander larva and an axolotl larva and found that axolotl’s
pituitary gland is as efficient as that of salamander larva. But the tissues of
axolotl failed to respond to the pituitary gland of the salamander. This
indicates the irrespon​sive nature of axolotl’s tissues to hormones.
● The investigations carried on by Etkin (1968) indicated that ‘spacing’ of
events during metamorphosis depends on thyroxin-concentration, while
the ‘sequence’ of events is inherent in the larval tissues. In amphibian
development the tadpole larva undergoes progressive metamorphosis and
transforms into an adult. This is a normal occurrence in amphibians.

But deviation from the normal pathway of development is found in the life-cycle of
many urodeles. Such devi​ated pathway of development in axolotls due to
extrinsic as well as intrinsic environmental factors may be regarded as
‘canalisation’, i.e., buffering of development against environ​mental change.

4. Genetics of Neoteny:
The role of thyroxine in urodele meta​morphosis has been revealed by studies in
genetics. The thyroid hormone binds to nucle​ar receptors that are in immediate
contact with DNA. The hormone helps to change the transcription of genes that
also influence to develop the larval characteristics to one, these gradually change
into juvenile and adult characteristics.
But the exact role of genetics in paedo​morphosis or neoteny has not clearly
under​stood. However, a single gene hypothesis is thought to control the axolotl’s
life cycle. 1: 1 Mendelian segregation results of paedomorph or metamorphic
backcrosses between A. mexi​canum and A. tigrinum.

If this theory is correct, then there are two alternate alleles at a major locus
primarily responsible for deter​mining the expression of metamorphosis or
paedomorphosis.

5. Significance of Neoteny:
● Neoteny is looked upon as a ​conse​quence of adaptation​ to neighbouring
envi​ronments where retention of larval gills and other larval features may
be advantageous.
● Weissmann (1875) regarded neoteny as a case of reversion to atavistic
ancestral conditions by assuming that all amphibia were originally
gill-breathing aquatic creatures and that every feature seen in a larva
represented a phylogenetic stage and the axolotl as such is a case of
reversion to an ancestral stage.
● External gills of urodeles have been evolved as an adapta​tion to aquatic
life. The external gills actually developed first in the embryos as additional
respiratory organs. The external gills were first initially restricted to
embryonic life, which may be prolonged in aquatic larval life.
● Possession of long external gills in the viviparous embryos of Salamandra
altra sup​ports the contention that the external gills are embryonic but not
larval features. So exis​tence of such gills in neotenous larvae is a
secondary but not an atavistic feature. Besides external gills, the tail with
tail-fin and epider​mal sense organs of the neotenous larvae are secondary
acquisitions rather than ancestral reminiscences.
● G. K. Noble (1931) regarded that the retention of larval features during
sexual matu​rity has nothing to do in the phylogeny of the amphibians. This
is quite evident from the heterogeneous characters of the
Perennibranchiata where all the neotenous species are included. Neoteny
as such may have some importance in the individual groups.

2. Types of Neoteny:
Kollmann (1882) classified neoteny into two categories, viz., Partial neoteny and
Total neoteny.

i. Partial neoteny:

Partial neoteny involves the simple postponement of metamorphosis beyond the

normal period. Retardation of metamorphosis may be due to temporary change
in ecological condition or due to sudden physiological abnormality.

ADVERTISEMENTS:

Wintering of the tadpdles of Pelobates fuscus, Pelodytes punctatus, Alytes

obstetricans, Hyla arborea, Bufo vulgaris, Bufo viridis, Rana temporaria, Rana
esculenta, Bombinator pachypus and many others furnish the typical examples of
partial neoteny.

In Alytes the broods usually complete their development within autumn. But the
larvae which hatched later in the months of July or August usually hibernate and
retain their larval features up to the next autumn.

In Rana esculenta most of the larvae attain much larger size than the normal and
remain in an under-developed stage, i.e., the hind limbs reach the budding stage,
the head and the body attains a blotted and unhealthy look. Majority of the larvae
remain in this slug​gish stage for one or two years after which metamorphosis
takes place.

ii. Total neoteny:

ADVERTISEMENTS:
In this category the speci​mens become sexually mature at the larval stage but
retain larval characters, like (i) exter​nal gills, (ii) tail-fin, (iii) ill-developed eyes, (iv)
ill-developed fin on the back and (v) very weak limbs. Total neotenic animals are
paedo-genic.

Because paedogenesis involves the capability of reproduction at the larval stage.

In this case of total neoteny the sexually functional larvae cease to
metamorphosis. In extreme cases of total neoteny (e.g., Necturus, Proteus, etc.)
the larvae attain sexual maturity and they remain in that stage without
undergoing metamorphosis.

Intermediate stages between partial and total neoteny are also recorded where
the lar​vae become sexually functional and may metamorphose into adults with
the advent of favourable conditions. The sexually mature axolotl larvae come
under this category.

Total neoteny is seen in many urodele (e.g., Necturus, Amphiuma, Triturus

vulgaris, Triton cristatus, Triton waltli, Ambystoma, Triturus alpestris, Siren,
Proteus, etc.). The phe​nomenon of neoteny has been extensively studied in case
of Ambystoma which, however, does not show an extreme case of neoteny. The
axolotl larvae of Ambystoma can metamor​phose into adult Ambystoma under
favourable conditions.

ADVERTISEMENTS:

Each axolotl larva possesses three pairs of delicate bushy external gills, four
pairs of gill-slits, a flat long tail with prominent tail- fin and a dorsal fin merged
with tail-fin. The axolotl larvae possess the power of regenera​tion. Chauvin has
experimentally shown that both accidental and experimental damaging of the gill
is followed by quick healing without affecting the process of metamorphosis.

The axolotl larvae become sexually mature when they attain the age of 6 months
only. It has been recorded by Metzdorff that the sexu​ally mature axolotls breed
during December or April to June. The spermatopores deposited by the male
members are drawn by the females into their cloacal chambers in the following
night.

On the following day, each female axolotl collects a leaf of aquatic plant and
pushes it against her vent. A few packets of eggs (each packet containing 6-10
eggs) are expelled violently into the surrounding water. Such violent expulsion of
the eggs is caused by the twisting and turning of the body of the female.

About 30 eggs are normally laid at a time. After a rest for a considerable period,
the female axolotls again start lying the eggs. The new-born larvae are hatched
out within 15 days. The optimum temperature for the deve​lopment of the eggs is
18-24°C. Bedriaga has recorded that the larvae at first feed on Daphnia,
Infusoria, etc.

ADVERTISEMENTS:

At the age of 6 months, the axolotls attain a length of 2.0-2.5 cm and become
sexually mature to start breeding. At this stage, the axolotls take up Tubifex,
scraped meat and may even scramble with the external gills of other fellows. The
damaged gills, how​ever, are quickly regenerated.

3. Factors for Neoteny:

The significance and cause of neoteny in Amphibians are not properly
understood. Several extrinsic and intrinsic factors are con​sidered to be
responsible for such unusual phenomenon.

Extrinsic factors influencing neoteny:

Gadow (1901) advanced the idea that the cause of retention of larval features in
axolotl is the abundance of food and other favourable requisites in aquatic life.
Shufeldt holds that deep water and cold​ness inhibit thyroxine secretion which
retards metamorphosis.
Drying up of swamps, lack of adequate food supply and rise in temperature in
surrounding water induce metamorphosis. Weissmann again claimed that the
retardation of metamorphosis of the axolotls is possibly due to the saline nature
of the water of the lakes where they live.

In an investigation to establish the role of physical factors in neoteny, tadpoles

were kept in water-holes with high vertical walls, so that they were not allowed to
reach the land above the water-holes. It has been observed that this forced and
prolonged use of larval gills and tails cause their further development, whereas
the growth of limbs and other structures nec​essary for terrestrial life remained
suspended.

Marie Von Chauvin in the University of Freeburg undertook similar experiments

with axolotl larvae and also came to the above con​clusion. It has further been
observed that the axolotls which were not likely to metamor​phose in normal
habitat could be forced to metamorphose by slowly accustoming them to
land-life.

Reverse phenomenon also happened in the life of axolotls. Partly

metamorphosed axolotls accustomed to land-life could again the induced to
return to larval life. All these experimental evidences emphasised the effect of
physical factors on metamorphosis.

Huxley (1929) undertook an investigation on temperature-coefficient on

metamorphosis. Several half-grown anuran tadpoles were cul​tured in variable
temperatures ranging from 3-30°C for same span of time. But the culture solution
has same concentration of thyroxine.

The larvae exposed to temperature range below 5°C could not complete their
metamor​phosis even when exposed to higher tempe​rature. But the larvae
exposed to higher temperature from the beginning completed metamorphosis
very quickly.

Despite extensive researches on the role of extrinsic factors on metamorphosis,

there is no sound reason to believe the exclusive role of the extrinsic factors. It is
commonly observed that the neotenic as well as normal over-grown larvae occur
side by side in the same habitat having similar environment.

So the existence of other factors besides the extrinsic factors becomes apparent.
There are internal and physiological factors which control neoteny. But the
extrinsic or environmental factors exert influence on the internal or intrinsic
factors.

Intrinsic factors influencing neoteny:

Many experimental evidences have been advanced by different investigators.

Zondeck and Leiter (1923) established that calcium delays metamorphosis in
axolotls. Gressner (1928) also advanced that insulin hormone inhibits
metamorphosis.

But recent researches incline to reveal that the metamorphosis is primarily

influenced (i) by varying threshold levels of thyroxine and its analogs and (ii) by
the degree of responsiveness of the larval tissue to hormones.

During early pre-metamorphic stage in amphibian development, the level of

thyroxine (T4) is kept very low in the body by genetic mechanism. Etkin and his
colla​borators have also established the role of pro​lactin on metamorphosis.

They have shown that the level of prolactin which acts as an inhibitor in the
overall control of metamorpho​sis remains high at this time. In the light of modern
genetics it may be suggested that the structural genes guiding the synthesis of
thy​roxine are ‘switched off’ by some operator genes whereas the genes guiding
the formation of prolactin are ‘switched on’.

ADVERTISEMENTS:

In such condi​tion the hypothalamus becomes sensitive to the available level of

thyroid hormone in the blood stream. The neurosecretory apparatus of the
hypothalamus produces a substance, called thyrotropin-releasing factor (TRF).
TRF stimu​lates the anterior lobe of pituitary to produce thyroid-stimulating
hormone (TSH) which in turn enhances the rate of thyroid secretion.

As the level of TSH rises during pro-metamorphosis, the level of prolactin

suddenly falls. So the metamorphosis starts. The time of shift in hor​mone
balance is possibly determined by the initiation of positive thyroid feed-back to
the hypothalamus. Poor secretion of thyroid glands and the irresponsiveness of
the larval tissues to the hormone are responsible for neoteny.

Kuhn (1925) studied the thyroid glands of neotenous larvae of the warty newt
and observed that the alveoli which secrete thy​roxine remain in undeveloped
state (only about half the size of those in normal full- fledged specimens). He has
further noted that the axolotls possessing normally developed thyroid glands
failed to pour their secretion in the blood stream.

Transplantation of at least two more thyroid glands in addition to the nor​mal

glands causes the metamorphosis of axolotls. This result indicated that the
thyroid gland of axolotls is able to produce one-third of the required quantity of
thyroxine and that the larval tissues are only one-third responsive in contrast to
normal specimens.

Bytenski and Saez have experimentally exchanged the pituitary gland between a
sala​mander larva and an axolotl larva and found that axolotl’s pituitary gland is
as efficient as that of salamander larva. But the tissues of axolotl failed to
respond to the pituitary gland of the salamander. This indicates the irrespon​sive
nature of axolotl’s tissues to hormones.

The investigations carried on by Etkin (1968) indicated that ‘spacing’ of events

during metamorphosis depends on thyroxin-concentration, while the ‘sequence’
of events is inherent in the larval tissues. In amphibian development the tadpole
larva undergoes progressive metamorphosis and transforms into an adult. This is
a normal occurrence in amphibians.

But deviation from the normal pathway of development is found in the life-cycle of
many urodeles. Such devi​ated pathway of development in axolotls due to
extrinsic as well as intrinsic environmental factors may be regarded as
‘canalisation’, i.e., buffering of development against environ​mental change.

4. Genetics of Neoteny:
The role of thyroxine in urodele meta​morphosis has been revealed by studies in
genetics. The thyroid hormone binds to nucle​ar receptors that are in immediate
contact with DNA. The hormone helps to change the transcription of genes that
also influence to develop the larval characteristics to one, these gradually change
into juvenile and adult characteristics.

But the exact role of genetics in paedo​morphosis or neoteny has not clearly
under​stood. However, a single gene hypothesis is thought to control the axolotl’s
life cycle. 1: 1 Mendelian segregation results of paedomorph or metamorphic
backcrosses between A. mexi​canum and A. tigrinum.

If this theory is correct, then there are two alternate alleles at a major locus
primarily responsible for deter​mining the expression of metamorphosis or
paedomorphosis.

5. Significance of Neoteny:
Neoteny is looked upon as a conse​quence of adaptation to neighbouring
envi​ronments where retention of larval gills and other larval features may be
advantageous.

Weissmann (1875) regarded neoteny as a case of reversion to atavistic ancestral

conditions by assuming that all amphibia were originally gill-breathing aquatic
creatures and that every feature seen in a larva represented a phylogenetic
stage and the axolotl as such is a case of reversion to an ancestral stage.

External gills of urodeles have been evolved as an adapta​tion to aquatic life. The
external gills actually developed first in the embryos as additional respiratory
organs. The external gills were first initially restricted to embryonic life, which
may be prolonged in aquatic larval life.

Possession of long external gills in the viviparous embryos of Salamandra altra

sup​ports the contention that the external gills are embryonic but not larval
features. So exis​tence of such gills in neotenous larvae is a secondary but not an
atavistic feature. Besides external gills, the tail with tail-fin and epider​mal sense
organs of the neotenous larvae are secondary acquisitions rather than ancestral
reminiscences.

G. K. Noble (1931) regarded that the retention of larval features during sexual
matu​rity has nothing to do in the phylogeny of the amphibians. This is quite
evident from the heterogeneous characters of the Perennibranchiata where all
the neotenous species are included. Neoteny as such may have some
importance in the individual groups.