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Water Vascular System of Echinoderms

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121 views

Water Vascular System of Echinoderms

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
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Water Vascular System of

Echinoderms
In this article we will discuss about Water Vascular
System of Echinoderms:- 1. Introduction to Water
Vascular System 2. Contents of Water Vascular System
3. General Plan 4. Modifications 5. Functions.
Introduction to Water Vascular System:
The water vascular system is enterocoelic in origin and
arises from the left hydrocoel. It exhibits radial symmetry
from the beginning and is equally developed in all
Echinoderms.

This system lies just above the haemal system. It is primarily


locomotory in function and also sub-serves the function of
tactile and respiratory organs in some cases. The excretory
role of water vascular system, suggested by some workers, is
not yet fully ascertained.

ADVERTISEMENTS:

Histological picture reveals that the canals have an inner


lining of flat ciliated epithelium, a layer of longitudinal
muscles, a connective tissue layer and an outermost layer of
flat ciliated cells.

Contents of Water Vascular System:


The canals of the water vascular system contain a fluid of
albuminous nature. It contains sea water and leucocytes.
Existence of red corpuscles is recorded in an Ophiuroid,
Ophiactis virens. Binyon (1964) has shown that the level of
potassium in the fluid may be as much as 60% above the sea
water value. Boolootian (1966) has recognised 14 different
types of amoebocytes in this fluid.

General Plan of Water Vascular System:


The water vascular system in different classes of
Echinodermata has almost the same structural organisation.
It comprises of a few canals together with some appendages
attached to these canals. The typical arrangement of the
water vascular system is exhibited by Asterias.

The water vascular system includes a circumoral canal


(circular ambulacral or ring canal) situated around the
mouth which gives tubular radial extensions, called radial
canals. The number of the radial canals is usually five. But
the number corresponds to the number of the radii of the
body.

ADVERTISEMENTS:

Each radial canal ends blindly at the end of the arm and
gives off along its course lateral vessels, each joining a tube-
foot. Each tube- foot is a hollow conical or cylindrical
process with an ampulla and a terminal sucker. The junction
between the lateral vessels and the tube-feet is provided
with valves which assist in locomotion.

The contraction of the ampullae results in the extension of


the tube-feet. A short, slightly curved, cylindrical and verti-
cally disposed stone or sand canal is present between the
madreporite and the ring canal. The stone canal opens into
the ring canal at the oral end and into the madreporic
ampulla at the aboral end.

The madreporite is a skeletal plate-like structure placed at


the aboral side. It is perforated by pores, called the
madreporic pores, which lead into madreporic ampulla or
vesicle from where the stone canal starts. The stone canal is
surrounded by a wider canal, called axial sinus, the wall of
which becomes folded to form the axial organ or dorsal
organ or ovoid gland or heart. The role of axial organ is not
fully known.
Besides the main vessels, some appendages become
associated with the system. Inter-radially located and
connected with the ring canal, there are polian vesicles and
Tiedemann’s bodies. The Polian vesicles are bladder-like
sacs with narrower neck.

They are contractile and usually manufacture amoeboid


cells. The Tiedemann’s bodies are glandular in nature and
consist of a number of branched tubules. They are yellowish
in colour and give origin to cells for the water vascular
system.

Modifications of the Water Vascular System in


Different Classes:
The water vascular system is equally developed in all
Echinoderms and has basically the same structural plan. In
the different classes, slight deviations from the basic plan
are encountered. The variations are due to their adaptations
to different modes of living.

Madreporite:
In Asteroidea (Fig. 21.7B), it is a calcareous sieve-like plate
and is situated aborally. The increase in number of the
madreporite is observed in many Asteroidea. The number of
madreporites is 3 in Asterias capensis, 4 in A, tenuispina, 16
in Acanthaster echinites. The madreporite is provided with
many secondary water-pores. Most of the water-pores lead
into stone canal and rest into the axial sinus in adults.
The water-pores are many in number and develop from one
primary larval water-pore. Like Asteroidea, in Echinoidea
(Fig. 21.16) also the madreporite possesses many pores, but
Echinocyamus pusillus, is peculiar in having only one water-
pore. In Ophiuroidea, the madreporite has one water-pore,
but in Ophiurae and Astrophytidae there are several water
pores.

In Holothuroidea true madreporite is absent. Great


variations are observed regarding the opening of the stone
canal. In Pelagothuria it opens to the exterior by one pore
and in many Elasipodidae there are 2 to 50 or more pores.
But in some Elasipodidae and Molpadidae the stone canal
opens into the coelom by many pores instead of opening to
the exterior.

In the rest of the Holothurians, the stone canal opens into


the axial sinus which in turn opens to the exterior by one or
more water-pores which are comparable to madreporite. The
madreporite in this case may best be called as internal
madreporite.
In Crinoidea, madreporite is represented by fine water-pores
on the body surface and these water-pores lead directly into
the body cavity. The water-pores are recorded to be 1500 in
Antedon bifidia.

ADVERTISEMENTS:

Stone canal:
Normally the stone canal is a short, slightly curved and
vertically disposed cylindrical tube. It opens into the ring
canal at the oral end. It is enclosed by the wall of another
wide canal, the axial sinus.

In Asteroidea, the stone canal is one and ‘S’-shaped. But in


Asterias rubens, there are two stone canals. The wall of the
stone canal is provided with calcareous ossicles. Devel-
opment of a longitudinal ridge-like projection makes the
stone canal complicated in the different members of the
Asteroidea (Fig. 21.37).
The following conditions are encountered:
ADVERTISEMENTS:

(1) In Echinaster purpureus, the fold projects as a ridge into


the canal. This represents the simplest condition.

(2) In Asterina gibbosa, the free terminal end divides into


two lamellae which may be coiled. This is seen in Asterias
and Gymnasterias.

(3) In Astropecten, the coiled lamellae become very


complicated and extend between the walls from one side to
another of the lumen.

ADVERTISEMENTS:
(4) In Culcita and Astropecten aurantiacus, the whole lumen
becomes divided into a number of irregular chambers.

In Echinoidea, the stone canal is only one and has soft


membranous wall devoid of calcareous matter. In Cedaris,
the wall of the stone canal is provided with calcareous
deposit. The stone canal has an ampulla below the
madreporite.

In Ophiuroidea, the stone canal is devoid of calcareous


deposition and opens in one of the oral plates (Sedjwick,
1898). In Trichaster elegans, there are five stone canals. In
Ophiactis virens, the stone canals are many.

In Holothuroidea, the stone canal is mostly single but in


some cases it may be more than one. The number of
accessory stone canal is also variable. Its walls are provided
with calcareous matters.

The opening of the stone canal shows greatest variation,


particularly in Holothurians. The stone canals in all
Holothurians are attached to body wall. In Pelagothuria, the
stone canal opens to the exterior by one or many pores. This
also is true in many Elasipodidae.

In Thy one, the stone canal is branched. In some


Elasipodidae and Molpadidae the stone canal ends blindly
and opens internally into the coelom by many pores as in the
genus Elasipoda.

In Crinoidea, stone canal as such is absent. Numerous tubes,


without calcareous deposits in their walls, emerging from
the ring vessel are the representatives of the stone canals of
other groups.

Axial sinus and axial organ:


The axial sinus is variously developed in different
Echinoderms. It is quite distinct from the perivisceral cavity
in adult excepting some Holothurians and Crinoids. The axial
sinus is inconspicuous in Asteroids, very small in Echinoids
and Ophiuroids. The axial organ, a fold from the wall of the
axial sinus, is present in all Echinoderms excepting
Holothurians.

The axial organ comprises connective tissue and cells of


germinal rudiment. In Echinoids the axial sinus ends blindly
and communicates with the stone canal. In Crinoids, the
portion of the coelom, into which the tubes from the ring
vessel open, represents the axial sinus. The axial organ
occupies the axis of the body. It consists of anastomosing
canals embedded in connective tissue.

Ring canal and Radial canals. The ring canal is a constant


structure in all Echinoderms and is situated round the
mouth. It gives tubular prolongations along the radii, called
radial canals or radial vessels. In Asteroidea, the ring canal
is pentagonal and is situated in the buccal membrane
(peristome). It is communicated with the exterior through
the stone canal and axial sinus.

In Echinoidea, the ring canal is situated at the upper end of


the jaws and gives five radial vessels. In Ophiuroidea, the
condition is same as in Asteroidea. In Holothuroidea, the
ring canal is situated around the oesophagus and the five
raidal vessels extend towards the oral end and again
proceed aborally along the radii of the body.

The radial vessels end blindly and the terminal tentacle,


characteristic of Asteroidea and Echinoidea, is absent. The
numbers of radial vessels are five. They are absent in
Synaptidae. In case of Crinoidea, the terminal tentacles are
absent and the radial vessels end blindly.
Lateral vessels and Tube-feet:
The radial vessels give lateral vessels to the tube-feet. The
tube-feet are cylindrical processes and their cavities are
continuous with the water vascular system. The tube-feet
possess ampullae at their inner ends and suckers at the
terminal ends. The ampullae are present in all echinoderms,
except Ophiuroidea and Crinoidea.

In Crinoidea, terminal suckers are absent and the tube-feet


are sensory and respiratory in function. In many
Astropectinidae, each tube-foot is provided with two
ampullae. In all the members of the Asteroidea the tube-feet
are provided with well-developed suctorial disc-like expan-
sions.

In Echinoidea, the tube-feet show variations. In Endocyclica,


the terminal ends of the tube-feet are suctorial and
supported by calcareous rings. In Cidaridae and
Echinothuridae, small oral tube-feet project from the
perforations of the ambulacral plates which are olfactory in
nature. In Clypeasteroids, the tube-feet are broad and the
walls are devoid of calcareous bodies. They help in
respiration.

The cylindrical tube- feet which are suctorial and provided


with calcareous rings, are locomotory in function. But in
Spatangoids, the tube-feet vary quite greatly which are due
to their functional activities.

The tube-feet without suckers are respiratory in function;


with suckers and calcareous ring are locomotory in function;
with expanded terminal disc and filaments around the mouth
as the tactile organ; rosette feet act as prehensile organs
and seize food from the surroundings.
In Ophiuroidea, the orientation of the lateral vessels and the
tube-feet is same as in Asteroidea, but they are devoid of
ampullae and are exclusively sensory in function.

In Holothuroidea, lateral branches from the radial vessels go


into the tube-feet as well as into the tentacles. Some lateral
branches also emerge from the radial vessels and end blindly
in the body wall. Ampullae are present in the tube-feet and
in the tentacular canals. The tentacular canals are devoid of
ampullae in Elasipodidae where they arise directly from the
ring canal.

Among the Crinoidea, in Antedon, each lateral branch from


the radial vessel supplies three tube-feet. The tube-feet have
ampullae. They are purely respiratory and sensory in
function.

Polian Vesicle and Tiedemann’s Bodies:


The ring canal possesses bladder-like polian vesicles and
gland-like Tiedemann’s bodies. In Asteroidea, the number of
polian vesicles varies greatly. They are totally absent in
Asterias rubens and A. glacialis. There are cases where two
or many polian vesicles may be present in each inter-radius
as seen in Astropecten.

In this case, a few vesicles open into the ring canal by one
common stalk (Fig. 21.38). The Tiedemann’s bodies are
attached to the ring canal and are usually two in each inter-
radius excepting that containing the madreporite where only
one is present.
Amongst Echinoidea, in most Endocylica, a small spongy
outgrowth in each inter-radius is present which is supposed
to be the polian vesicle. There are five Tiedemann’s bodies in
Echinoidea. In Ophiuroidea, in each inter-radius excepting
that of stone canal, there is a polian vesicle.

In Ophiactis virens, besides two or three polian vesicles


opening in each inter-radius, there are many tubular canal of
Simroth (supposed to be respiratory in function). The
Tiedmann’s bodies appear to be wanting. Some authorities
refer some structures homologous with Tiedmann’s bodies.

Some say that the radiant protrusions are found in some


places. Hyman (1955) also refers that these are Tiedmann’s
bodies. Fedetov (1926) has reported that the radial
protrusion is associated with water ring in Ophiactum
sericeum.

In Gorgonocephalus, a bunch of pouch-like structures or


branching tubules are present in water ring. He did not
mention it as Tiedmann’s bodies but represented it as spe-
cialized structure.
Hyman (1955) said perhaps this structure mentioned in
these above animals as homologous with the Tiedmann’s
bodies. In Holothuroidea, usually one large polian vesicle is
present. In some exceptional cases more than one polian
vesicle may be present. In Crinoidea, the polian vesicle and
Tiedemann’s bodies are absent.

Functions of the Water Vascular System:


1. Locomotion:
The main function of the water vascular system is to help in
locomotion. Echinoderms having suctorial podia (tube-feet)
can adhere to the substratum temporarily. The mechanism
of locomotion has discussed in detail under the water
vascular system of Asterias and Echinus.

2. Respiratory and sensory:


In Ophiuroidea and Holothuridea the tube-feet (podia) are
primarily sensory in function. In Echinoidea (in regular
urchins), the tube-feet of the aboral side lack terminal disc
and are sensory in function (Hyman, 1955).

In spatangoids, the petaloids of the aboral surface are


provided with lobulated podia without suckers and are
believed to some respiratory in function (Loven, 1883).

3. Excretory:
Nitrogenous wastes are eliminated through the thin areas of
the body surface such as the walls of tube-feet.

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