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Evolution of Tetrapod Limbs in Amphibians

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12 views7 pages

Evolution of Tetrapod Limbs in Amphibians

Uploaded by

apranav2005a
Copyright
© All Rights Reserved
We take content rights seriously. If you suspect this is your content, claim it here.
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Download as DOCX, PDF, TXT or read online on Scribd

The evolution of the tetrapod limb was a

significant hallmark in vertebrate evolution.


The first tetrapods were amphibians that
evolved from the crossopterygian fishes that
lived in shallow marshy locations and already
possessed lungs for respiration and lobed fins
to support their bodies on muddy banks of
ponds, rivers and marshland.

Amphibian Origin
They are the first vertebrate to walk on land.
Recent mtDNA evidence shows that modern
amphibians are more similar to lungfishes
(Dipnoi); Plenty of selective pressure for
amphibians to emerge onto land - Wet and
dry spells; competition in aquatic; Oxygen
low waters; Tremendous radiation of
arthropods in the terrestrial environment;
Ability to move out of water escaping from
aquatic predators.
Amphibians are distributed in water and land
throughout the world, except in Antarctica and Greenland, where there are no amphibians. There are about
7000 different species of amphibians.
The anatomical changes that took place during the transformation from fishes to Amphibia were as
follows:
 A tetrapod limb evolved to support the
body out of water as on land
 Gills were lost in the adult stage as lungs
became more and more efficient air-
breathing organs
 The hyomandibular bone of the second
visceral arch transformed into columella
to transmit sound vibrations from air to
the inner ear.
 Loss of scales permitted cutaneous
respiration, which evolved as an
alternative method of respiration in the
absence of gills while the animal lived
and swam in the water.
 The skull developed two occipital condyles that fitted in an atlas vertebra so that the skull could move
upward and downward for locomotion and better visibility.
 In the terrestrial environment, the vertebral column must be strong to support the body weight, and hence
centra of vertebrae transformed from amphicoelous to procoelous type.
 The lateral line system that was not useful in the terrestrial environment disappeared.
 Adipose tissue stores fat and provides insulation developed under the skin to stop water loss by
evaporation on land.
(Amphicoelous: Most teleost fish, a few salamanders; centrum hollowed at the anterior and posterior end.
Procoelous: Anurans & present-day reptiles; centrum concave on the anterior surface, convex on posterior
surface).

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Amphibia General characters:
• Amphibians are cold-blooded vertebrates that can live on land and in water- i.e., amphibious. None are
true marine forms
• The body divisible into head and trunk; Tail is present in urodela animals.
• Amphibians show four limbs, may vary in size, with the forelimbs of some being much smaller than the
hind with which they can swim in the water and jump or walk on the land, except apoda
• Limbs have varying numbers of digits, and webbed feet are often present; no true nails or claws.
• limbs evolved from the pectoral and pelvic fins
• The eyes developed lids and associated glands, which aid the protection of eyes terrestrial environment.
• The exoskeleton is absent (apoda animal's small cycloid scales are present); the skin is smooth, thin,
hairless, and porous; it contains both mucous glands and poison glands.
• A bone endoskeleton with varying numbers of vertebrae;
• Respiration is accomplished either separately or in combination by lungs, skin, and gills; some larval
types possess external gills, which may persist throughout life.
• The Amphibian's skull is dicondylic.
• The mouth is usually large with small teeth in upper or both jaws; two nostrils open into the anterior part
of the mouth cavity.
• Amphibian dentition consists of an upper crown and a lower base (pedicle). However, teeth are generally
not used for chewing but for holding the prey firmly.
• The digestive system is well developed. A well-developed liver is present
• The external ear is absent. They have external tympanum (eardrums); Middle and inner ears are present;
• Amphibians Heart is 3 chambered with 2 auricles and 1 ventricle; blood contains R.B.C; they are
nucleated; contain hemoglobin; a pulmonary and a systemic circuit
• The blood vascular system contains hepatic and renal portal systems.
• Amphibians Kidneys are mesonephric; Urinary bladder is present
• Well-developed Central nervous system; the brain occupies the cranial cavity completely. The brain is
divided into fore, mid and hindbrains. Brain continuous as spinal cord; 10 pairs of cranial nerves
• Sexes are separate, Sexual dimorphism; internal or external fertilization; a larval stage may be present;
egg does not have a shell; Eggs are telolecithal, holoblastic cleavage (entire cleavage) & unequal
• In a few forms of amphibians, the adults will retain larval characteristics = neoteny. Ex: axolotl larva
• Amphibians have a vocal sac that resonates and amplifies the vibrations from the vocal cord.
• Chromatophores: highly branched asymmetrical shape cells with color pigments in them.
Xanthophores: red to yellow color; xanthophylls; Melanopores: dark brown & black; melanin;
Iridophores: whitish reflective granules; guanine
• Metachrosis: change in color intensity of skin to blend with the external environment. Changes are
depending upon the temp and moisture, avoiding detection by enemies/prey. Dry and bright light
condition: chromatophores contract and their pigments granules aggregates in the middle, skin color
lightens. Humid and dim light: Chromatophores expand & granules disperse, skin becomes darker.

Classification of Amphibians
 Subclass Labyrinthodontia –Earliest ancestors with labyrinthine folds in teeth. Extinct.
 Subclass Lepospondyli –With scale-like ossification in vertebrae. Ancestors of caecilians. Extinct.
 Subclass Phyllospondyli - Ancestors of urodeles and anurans with leaf-like ossification around the
notochord. They had short body and tail. Extinct.
 Subclass Lissamphibia – frogs, toads, salamanders, newts, etc. Modern living amphibians.
• Order Anura (frogs and toads)- lack tails as adults
• Order Urodela/Caudata (salamanders, newts, sirens, etc.)-retain tails as adults
• Order Gymnophiona /Apoda (Caecilians)-lack legs

Order ANURA or SALIENTIA: Frogs and Toads. Total 5453 species. Frogs and toads are tail-less; they
have large, well-developed eyes, short, thick bodies and four legs. Frogs have long, powerful hind legs that
are well adapted for leaping and swimming. Tree frogs have suction pads on their fingers and toes for
climbing. Toads have shorter legs than frogs adapted for hopping or walking instead of leaping. The skin of

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toads has a dry, warty appearance. Indicator species: susceptible to environmental changes (E.g., UV
exposure and water pollution) due to dependence on the moist/ aquatic environment.
Order CAUDATA or URODELA Giant salamanders, Asiatic salamanders, Mudpuppies, newts, Sirens.
Tailed amphibians. Total 560 species. Salamanders have a tail, short bodies, four almost equal legs, and
moist skin. They are often brightly colored and have great powers of regeneration. Generally, 4 limbs at
right angles to the body, adapted to walking rather than jumping or hopping. Neoteny: Retention of juvenile
characteristics in the adults of a species, as among some species
Order GYMNOPHIONA or APODA or CAECILIA: limbless amphibians. Total 170 species; look like
worms or snakes with a short-pointed tail. They are well adapted for burrowing in moist sand. The long,
slender body has numerous grooves. The eyes are small and covered by skin. There is a tentacle that is a
tactile organ. Caecilian species range in length from about 10 cm to nearly 1.5 meters. Have fishlike scales.

Rana tigrina external morphology

Shape, size and color : Spindle shaped,


dorsoventrally flattened, streamlined- offer less
resistance to water in swimming and to hopping in
the air; Moderate size, 12 to 20 cm; Dorsal body
surface is spotted olive green; with dark bands;
Ventral pale yellow; The frog body may be divided into head and trunk.
Head: Triangular flat, contains the mouth, eyes, eardrum and nostrils; Snout blunt, wide mouth terminal;
Eyes have nictitating membranes which protect the eyes from particles while swimming; A pair of small
circular external nostrils, located dorsally at the tip of snout, open into the anterior part of buccal cavity; A
small, but intensely pigmented circular patch of tough, membrane-like skin, called tympanic
membrane or eardrum, is located behind each eye- receives sound waves.
Trunk: The compact trunk first widens and then tapers between the hind limbs; A short distance behind its
middle, it is dorsally raised into a sacral hump; At the posterior end, it has the median cloacal aperture;
common went for excretory and reproductive products; Two pairs of limbs are attached to it.
Limbs: Forelimbs are short and robust; Forelimb consist of the upper arm (brachium), a forearm
(antebrachium), and a hand (manus) with four digits; The first digit, i.e., the thumb (=pollex), is absent; The
forelimbs help in directional orientation during locomotion and bear the shock of body weight on landing
after a leap; Hind limbs very long and powerful folded in the manner of 'Z'; Hind limbs are long consists of
thigh, shank and a foot with five digits. All toes of a hind limb are joined together by a thin web of skinfold;
Hind limb adapted for both leaping and swimming; Digits of all limbs are without claws or nails.
Frog's skin is soft, smooth, and loose; It is always kept moist and slimy by mucus secreted by cutaneous
glands; Frog’s skin serves as an essential accessory respiratory organ, also helps the frog in slipping out of
the grip of an enemy; The dorsal skin shows a few low folds called dermal plicae.
Male& female: male larger, male possess vocal sacs; males possess a special
nuptial pad.

NEOTENY in Amphibians
Neoteny is defined as the failure or delay of the larva to metamorphose while
becoming sexually mature. It is the character of some amphibians. The best
example is axolotl larva of Amblystoma. It is aquatic. It has gills. It develops
gonads. It lays eggs and attains a large size. Amblystoma, on the other hand, is

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terrestrial and without gills. Proteus and Necturus are permanently neotenous forms. They retained the larval
features and reproduced sexually like a mature animal.

RESPIRATORY SYSTEM IN FROG


All frogs start life as aquatic tadpoles, breathing underwater through internal gills and their skin
Later most develop into land animals with lungs for breathing air
Tadpoles respire gills
Adult frogs lose the gills,
Gas exchange between the air and bloodstream occurs in three locations in the frog
 the lungs: pulmonary respiration
 the skin: cutaneous respiration
 the mouth; buccal breathing
Cutaneous respiration (30% of respiration)
The skin is useful for breathing on land or in water and is in use continuously in water;
hibernation/aestivation skin is the only organ of respiration; it has no exoskeleton; it is highly vascularized
with most of its good network of thin-walled capillaries located next to its thin epidermis; the mucus
secreted by its glands (cutaneous glands) always keeps its surface moist and always holds a thin film of
water between itself and the skin surface. Gaseous exchange occurs in a solution between the blood of
cutaneous capillaries and the outside environment (water or air) through a thin film of water. Underwater, or
when buried deep under mud during the winter and summer sleep (hibernation and aestivation), frogs can
naturally respire only by their skin. However, even during their stay on land in spring and rain, cutaneous
respiration continues as usual. Certain small amphibians, like the Plethodont salamanders, have no lungs
and respire only by their skin
Bucco- Pharyngeal Respiration (5% of total 02-intake)
Like the skin, the mucosa of the buccopharyngeal cavity in the frog is also ideal for gaseous exchange.
It occurs when frog quietly floating upon the water surface, and even when resting upon the land. The
mucous lining of the buccal cavity richly supplied with blood capillaries- remains moist by mucous- gas
exchange takes place readily. During this process, the mouth, gullet, and glottis remain closed, but nares
remain open. Atmospheric air is sucked into the cavity through nares when the floor lowers, and it is forced
out when the floor rises. The alternate lowering and rising of the buccal cavity; buccal respiration is bought
about
Oxygen dissolves in the mucous – gets into the blood. While at rest, this process is the predominant form of
breathing.
Pulmonary respiration (65% of the total 02-intake)
Comprises a pair of lungs and air passages leading to and from the lungs (a conducting
part & respiratory part). The conducting part includes nasal cavities, buccopharyngeal
cavity, larynx, trachea, etc. The respiratory portion includes the lung alveoli, where
gaseous exchange occurs between blood and air. The adult frog has very simple,
pinkish, sac-like organs called lungs, located anteriorly in the abdominal cavity on
either side of the heart. The lungs can be used only when the frog is on land or
swimming at the surface. The nostrils have valves that prevent water entry and control
the flow of air into the lungs. Unlike mammals, they do not make regular and rhythmic
breathing movements but gulp air into their lungs spasmodically as the need arises.
Both lungs lead in front of a small but vast common chamber, called bronchus.
Bronchus leads in front into another small and spherical box-like laryngotracheal
chamber , which opens into the buccopharyngeal cavity by the slit-like glottis. A
dense network of irregularly branched septa from the lung wall divides the lung cavity
into numerous small chambers called air sacs or alveoli. The alveoli wall is highly
vascularized; its internal lining is very thin, which always remains moist by mucus. Frog breath with their
mouth closed. Their throat movements pull air through the nostrils to the lungs. Then breathe out with body
contractions. Pathway of air: Nares, pharynx, glottis (opening to the trachea), trachea, bronchi, and alveoli
(air sacs). The exchange between air in the lungs and blood in the capillaries. A dense network of irregularly
branched septa, arising from the lung wall, divide the lung cavity into numerous small chambers called air

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sacs or alveoli. The alveoli wall is highly vascularized; its internal lining is very thin, which always remains
moist by mucus. Frog breath with their mouth closed
Their throat movements pull air through the nostrils to the lungs. Then breathe out with body contractions
Pathway of air: Nares, pharynx, glottis (opening to the trachea), trachea, bronchi, and alveoli (air sacs). The
exchange between the air in the lungs and blood in the capillaries.
Breathing occurs in four steps:
 The throat moves down, drawing air into the mouth through the nostrils
 The nostrils close and the throat moves up, forcing air into the lungs
 To eliminate the carbon dioxide in the lungs; the throat moves down drawing air back out of the
lungs
 The nostrils open and the throat moves up, forcing air out of the mouth

PARENTAL CARE IN THE MEMBERS OF CLASS AMPHIBIA


Parental care means care of the eggs or juveniles until they reach the reproductive age. Amphibians were the
first vertebrates to have evolved different kinds of parental care to protect their young ones. Parental care
evolved to reduce the energy expenditure on reproduction, as in the absence of it animals must produce
millions of eggs so that few could survive to replace the parents to ensure existence of the species.
Parents protect the eggs and early developmental stages in two ways.
1) They construct nests
2) Direct Nursing.
APODA (=GYMNOPHIONA)
The female Icthyophis glulinosa will dig a hole in the moist soil near a pond, deposit eggs in it and mother
will coil around egg mass protect from the enemies.
Mothers of the Kenyan caecilian, Boulengerula taitanus provide an outer layer of their modified skin as a
food source to their offspring.
URODELA (=CAUDATA)
a) Protection by Nests
Salamandrella keyserlingi will construct a gelatinous bag like structure attached to an aquatic plant below
the water, in which eggs are stored.
Autodax will lay eggs in a dry hole on the soil or in a hole. The parents also live in the hole and protect the
egg and the larvae developed from them.
b) Direct Nursing by Parents
Amphiuma, (Congoeel) The mother will coil around the eggs and protect them.
Desmognathus fuscus, (the dusky Salamander) the mother will carry the eggs attached around its neck.
Salamandra salandra, (the European fire salamander):- The mother will retain few eggs in its body and
liberates the larvae into the water, where they complete their development.
Salamandra atra, (the European alphine salamander): The mother keeps the eggs in its uterus. It liberates
only the young ones who are completed their metamorphosis.
ANURA:
In Aura amphibians, parental care is reached its peak.
a) Protection by Nests: Many frogs and toads build nests.
In these organisms the larva comes out in a very early embryonic stage which requires some protection in
the very early stages of development, hence the parent will build nests.
Hyla faber: - It is Brazilian free frog.
Rhacophorus schlegeli it is called Japan tree frog
Hyla resinfectris it is a free frog
Rhacophorus malabaricus - It lays eggs on the branches or leaves of a tree which will be hanging over a
pond. These larvae after hatching from eggs will fall into the pond water and undergo metamorphosis.
2) Direct Nursing by Parents: This method is more advanced than constructing nests.
Mid-wife-toad: female frog lay strings of eggs around males back and thighs. When the eggs are ready to
hatch the male frog moves to a nearby pond, and the larvae are released.

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Marsupial frog: It has a special pouch in its skin. It opens out through an opening near the cloaca. Fertilized
eggs are transferred into this pouch. The eggs are stored in this pouch where they ‘undergo development,
and tadpoles are liberated out.
Surinam toad (Pipa Americana): During breeding season the skin of the back of the female becomes soft
and spongy. During copulation, the oviducts will come out of the female- Because of the movements of
male, the eggs are forced out of the oviduct. Each egg sinks into a small pocket of the skin.
Little South American frog (Rhinoderma darwini): The fertilized eggs are transferred into its vocal sacs; the
development takes place. Then the completely developed young individual will jump out from the mouth of
the male frog.

GENERAL INFORMATION

TYPES OF EGGS
Different species of animal has a different type of eggs. They are classified on the following basis -
1. Based on the Amount of Yolk
(i) Alecithal - the yolk is almost absent. E.g., Eutherian eggs
(ii) Microlecithal - a minimal amount of yolk. E.g., Amphioxus and Tunicates.
(iii) Mesolecithal - moderate amount of yolk. E.g., Amphibian, Dipnoi Fishes, and Annelids
(iv) Mega/Macro/ Polylecithal - large amount.E.g. Insect, reptiles, cartilage fishes and Prototheria.
2. Based on the Distribution of Yolk
(i) Iso/ Homolecithal - even distribution of the yolk in the ooplasm. E.g., Alecithal and microlecithal yolk.
(ii) Telolecithal - unequally distributed, the yolk is found in the one pole which called as vegital pole. The
yolk free pole animal pole. E.g., fishes, amphibians, reptiles and birds. Yolk in the ooplasm.
(iii) Centrolecithal - yolk in the center and surrounded by ooplasm. E.g., Insect and hydrozoa.
3. Presence or Absence of Hard Shell
(i) Cleidoic eggs - eggs are covered by the hard shell for protection permeable for gases. Yolk present in
enough quantity. E.g., Reptiles and birds.
(ii) Non-cleidoc egg - They
are without shell and these
develop in aquatic medium
and uterus of female. E.g.
amphioxis, mammals, frog
and hardmania.
4. On The Basis of
Development
(i) Determinate or Mosaic
Eggs
In these types of eggs before
fertilization determined the
different parts of an embryo
E.g., Nimertine, Annelids,
Molluscs and Ascidian.
(ii) Indeterminate or
Regulative Eggs
Different parts determined
after the third cleavage; in
man twins developed due to
this region.

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DIFFERENT TYPES OF VERTEBRAE

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