Obelia: Habitat, Structure and

Diagram
Habit and Habitat of Obelia:

Obelia is sedentary, marine colonial form found attached on the

surface of sea weeds, molluscan shells, rocks and wooden piles in
shallow water up to 80 metres in depth. Obelia is cosmopolitan in
distribution, forming a whitish or light-brown plant-like fur in the
sea; hence, the common name sea-fur is assigned to it.

Obelia Colony – A Gross Structure:

Each colony of Obelia consists of a horizontal thread-like root

called hydrorhiza which is attached to a weed or any substratum.
From hydrorhiza arises a vertical branching stem about 2.5 cm
long which is known as a hydrocaulus. The hydrorhiza and
hydrocaulus are hollow tubes.

The hydrocaulus bears zooids or polyps on either side in a

cymose formation. At the growing ends of the main branches are
immature club-shaped polyps. Each polyp has a stem and a
terminal head called a hydranth. The hydranths are feeding
polyps, they feed by capturing minute animals and larvae.
Towards the base of the hydrocaulus in the axils of the polyps, are
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 reproductive polyps called blastostyles.

The polyps, their tubular connections and blastostyles are made

of ectoderm, mesogloea and endoderm, these layers are together
called coenosarc and its cavity is an enteron which is continuous
and common to all the members, through the enteron digested
food is distributed in solution.

The entire colony is covered by a tough, yellow chitin secreted by

the ectoderm, this covering is known as perisarc. The perisarc
constitutes the exoskeleton and it covers the hydrorhiza,
hydrocauli and their branches, and at the base of each polyp, it
forms a clear, wine glass-shaped hydro theca.

The hydrotheca has a shelf across the base which supports the
hydranth, and the hydranth can contract and withdraw into the
hydrotheca.

The perisarc around a blastostyle is a gonotheca, the blastostyle

and gonotheca are together called a gonangium. The perisarc is
an exoskeleton, at first it is continuous with the coenosarc but on
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 growing thick, it separates and is joined to the coenosarc only at
intervals by minute projections, at these places it gets ringed
which allows bending.

The Obelia is a trimorphic colony, that is, having three kinds

of zooids which are as follows:
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 1. Polyps or hydranths (nutritive zooids);

2. Gonangia or blastostyles (budding zooids);

3. Medusae, (sexual zooids).

In fact, to start with Obelia is a monomorphic form having polyp

only but later due to the development of blastostyle it becomes a
dimorphic colony and finally medusae bud over the blastostyle in
a mature colony, then it becomes a trimorphic colony.

Polyp or Hydranth:

The colony of Obelia has many polyps (Gr., polypus – many-

footed) or hydranths (Gr., hydra = water serpent; anthos = flower)
or gastro zooids. Each polyp is very much like a miniature Hydra.
It has a cylindrical body attached to the axis of the hydrocaulus by
its proximal end and free at its distal end. It is covered by a cup-
shaped hydro theca.

The free distal end is produced into a conical elevation, the

hypostome or manubrium which is about one-third of the length
of the hydranth. The hypostome is surrounded by a circle of
numerous (about 24) tentacles. The tentacles are longer than
hypostome, tapering and filiform.

The apex of the hypostome bears a terminal aperture called

mouth which is capable of great dilation and contraction.

Below the hypostome is the stomach region of the polyp. The

body and manubrium of the polyp enclose a spacious enteric
cavity or gastro vascular cavity. The polyp is protected in hydro
theca, which is prolongation of the perisarc. At the base of the
polyp, it forms ring-like horizontal shelf at which the polyp rests.
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 Histology of Polyp or Hydranth:

The polyps have an outer layer of ectoderm and an inner layer of

endoderm, between them is a thin, transparent mesogloea; all
these layers constitute the coenosarc which is soft and tubular,
the continuous cavity is the enteron or gastro vascular cavity.

The enteron has a fluid and its lining is flagellated. Rhythmical

contractions of the hydranths cause a current which distributes
food obtained by some polyps to those parts of the colony where
feeding is not taking place. The tentacles of polyps are solid with
no enteron. They have a single- layered core of vacuolated
endoderm cells with thick walls inside a layer of ectoderm.

Ectoderm:

It consists of long, conical columnar epitheliomuscular cells, their

inner ends are produced into muscular processes which run
longitudinally. In the ectoderm layer are very few interstitial cells,
some branching nerve cells and cnidoblasts with nematocysts.
The nematocysts are abundant on the tentacles and manubrium
only.

The cnidoblasts are found in the basal part of the hydranth and in
the coenosarc. They form nematocyst and migrate actively to
reach their final positions. Obelia has only one kind of
nematocysts called basitrichous isorhizas in which the capsule is
oval, butt is absent, the thread is open at the tip and has spines on
its base.
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 Endoderm:

It has long, granular epitheliomuscular cells, their muscle

processes point outwards and are circular. Endoderm cells have
flagella which produce a current in the enteron. They can also
form pseudopodia for engulfing food. The endoderm of tentacles
has cubical, vacuolated cells with thick walls.

In the endoderm layer are nerve cells and club-shaped gland cells
which produce digestive enzymes. Mesogloea is a thin jelly-like
substance with no structure or cells. On each side of the
mesogloea is a nerve net composed of nerve cells and their fibres,
the two nerve nets are inter-connected.

Polyp is the nutritive zooid of the colony. It is carnivorous and

feeds upon aquatic crustaceans, nematodes and other worms.
Tentacles help in catching and conveying the prey to the mouth.
Digestive juice is secreted in the gland cells of gastro dermis and
the process of digestion is extracellular as well as intracellular.

Gonangium:
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 The gonangium (Gr., gonos = seed; angeion = vessel) (Fig. 32.3)
is club-shaped, cylindrical form. It is covered by a transparent
gonotheca and contains an axis or blastostyle on which lateral
buds form that develop into medusae or gonophores. The
blastostyle has no mouth and no tentacles, but ends distally into a
flattened disc.

The gonotheca opens at its distal end by a gonopore, through

which the medusae escape. Gonotheca, blastostyle and the
gonophores together form a gonangium.

Asexual Reproduction:

When the temperature of the water exceeds 20°C, the buds

which would normally form gonangia in the colony break free from
the colony and settle down; a stolon arises from the lower end of
the bud which produces a new colony of Obelia asexually. This is
a special mode of asexual reproduction.

Medusa:

The medusa is a modified zooid produced as a hollow bud from

the coenosarc of the blastostyle in spring and summer. Medusa
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 swims freely on the surface water.

Structure of Medusa:

It is saucer-shaped, it is attached by the middle of the convex

surface to the blastostyle, when fully formed it breaks free and
emerges from the mouth of the gonotheca.

The medusa is circular and tiny umbrella-like in shape. The

convex outer surface is known as the ex-umbrella and the
concave inner surface is the sub-umbrella. From the centre of the
sub-umbrella arises a short projecting manubrium (L., manus =
handle), at its apex is a square mouth surrounded by four oral
lobes.

The mouth leads into an enteric cavity or gastric cavity in the

manubrium. From the enteric cavity, arise four radial canals which
are delicate ciliated tubes, they run to the margin of the bell to join
a ciliated circular canal running near the margin.

The enteric cavity and the canals represent the enteron which
distributes food. Projecting from the middle of the radial canals
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 are four gonads, since sexes are separate they are either four
testes or four ovaries,they are patches of modified sub-umbrellar
ectoderm.

The gonads mature after the medusae escape from the

gonotheca. The edge of the bell is produced inwards as a thin fold
called velum.

Velum is characteristic of hydrozoan medusae but it is

insignificant in Obelia. The medusae with a velum are called
craspedote, and those with no velum are acraspedote
(Scyphozoa). From the edge of the bell numerous small solid
tentacles hang downwards. The tentacles have swollen bases due
to the accumulation of interstitial cells which are practically
absent from other regions.

The basal swellings of tentacles are called vesicles or bulbs,

nematocysts are formed continuously in the bulbs from where
they migrate to the tentacles.

Digestive enzymes are secreted from the endoderm of bulbs.

Near the bulbs the ectoderm has pigment granules and nerve
cells, they are often called ocelli and it is claimed that the ocelli
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 are sensory to light, but more probably there are no ocelli, the
pigment granules are accumulated excretory matter.

Above the bulb of every tentacle is a tiny fluid-filled vesicle.

Nematocysts are confined to the manubrium and tentacles, there
may be some on the bell margin. There are eight marginal sense
organs called statocysts or lithocysts lying at regular intervals,
being attached on the sub-umbrella side to the bulbs of eight
tentacles, they are developed in response to a locomotory habit.

A statocyst is a tiny, circular closed vesicle lined with ectoderm

and filled with a fluid containing calcareous granules called
otoliths which lie in a special cell called lithocyte.

The lining has some sensory cells with thin sensory processes on
which the otoliths produce a stimulus which is transmitted by
nerves to muscles; the muscles coordinate the snake-like
swimming movements of the medusa, and should the medusa
become tilted, the muscles contract to right the position of
medusa bell, thus, statocysts are balancing organs.

Histology of Medusa:

The ectoderm covers the bell on all sides, its epitheliomuscular

cells are produced into muscle processes which run longitudinally
in the manubrium and tentacles. In the sub-umbrella, the muscle
processes of the ectoderm are so large in proportion to the
epithelial part that they almost form muscles only.

The muscle processes form a striated circular muscle and some

radial muscles in the sub-umbrella, they bring about locomotory
movements. The ectoderm of the ex-umbrella is devoid of
musculature.
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 The endoderm lines the enteric cavity and the radial and circular
canals. The endoderm cells have no muscle processes, they are
ciliated epithelial cells, they are digestive. Between the two
ectoderm layers of the bell is a thin sheet of endoderm lamella
except where the enteron lies.

The endoderm lamella is formed by the fusion of upper and lower

layers of endoderm, the fusion having occurred at all places
except in the region of the enteron. Between the ectoderm and
endoderm is thick mesogloea forming the bulk of the bell of the
medusa, manubrium and tentacles. The velum has a double layer
of ectoderm and the thick mesogloea in the middle, it has no
endoderm.
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 Radial symmetry of Medusa:

Like polyp, the medusa is radially symmetrical.

The presence of the four radial canals distinguishes the four

principal radii or per-radii. Halfway between any two per-radii a
radius of the second order or inter-radius may be taken. Halfway
between any per-radius and inter-radius on either side a radius of
third order, or ad-radius, and halfway between any ad-radius and
the adjacent per- or inter-radius, a radius of fourth order or sub-
radius.

Thus, there are four per-radii, four inter-radii, eight ad-radii and
sixteen sub-radii. In Obelia, the radial canals, the angles of the
mouth and four of the tentacles are the per-radial, four more
tentacles are inter- radial, and the remaining eight tentacles,
bearing the lithocysts are ad-radial. Sub-radii are of no
importance in this particular form.

Development of Medusa:

The blastostyle produces medusae by budding in large numbers.

The cavity of the blastostyle pushes the coenosarc out to form a
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 small protuberance or bud. The bud grows larger and its
coenosarc becomes like a vesicle which is attached to the
blastostyle by a narrow stalk. The cavity of the vesicle is
continuous with the enteron of blastostyle.

The distal ectoderm of the vesicle separates into two layers, then
the inner layer of ectoderm splits to acquire a cavity called a bell
rudiment. There are now two layers of ectoderm outside the bell
rudiment and one layer on the inner side. The cavity of the bell
rudiment assumes the shape of the sub-umbrella, and a
manubrium is formed in the centre.

The two layers of ectoderm which enclose the bell rudiment from
outside now break leaving a marginal and circular shelf called
velum.

In most hydrozoan medusae, the velum grows and becomes

prominent, but in Obelia it decreases and becomes insignificant.
The manubrium acquires a mouth, marginal tentacles are formed,
the stalk breaks and its aperture closes up, thus, a medusa is
formed which is set free, it escapes from the gonotheca, later its
gonads mature.
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 Nutrition:

Medusae are carnivorous and the processes of their nutrition are

essentially the same as in the polyp. The food consists of living
animals or bodies of animals.

Digestion is both extracellular and intracellular. Extracellular

digestion occurs in the main part of the gastro vascular cavity and
is purely proteolytic. Hyman (1940) has shown that although food
particles are distributed throughout the gastro vascular cavity,
most intracellular digestion takes place in the manubrium, in the
stomach and in tentacular bulbs.

The digested food is distributed to the entire medusa through the

system of radial and circular canals.

Muscular system:

The muscular system of medusa is somewhat more specialised

than in the polyp. The gastro dermal cells lack contractile
extensions, and the muscular system is, thus, restricted to the
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 epidermal layer. Furthermore, the muscular system is best
developed around the bell margin and sub umbrella surface where
the fibres form a radial and circular system.

Some of the epitheliomuscular cells of the velum have their

contractile extensions oriented to form a powerful circular band
of fibres which are striated. The contractions of the muscular
system, particularly of circular fibres produce pulsation of the
bell.

The swimming movement of the medusa is dependent base on

these pulsations and is largely vertical in lamella direction.
Horizontal movement is dependent upon water currents.

Nervous system:

The nervous system of medusa is more highly specialised than

that of the polyp. In the margin of the bell, the epidermal nerve
cells are usually organised and concentrated into two nerve rings,
one above and one below the attachment of the velum.

The nerve rings connect with fibres innervating the tentacles, the
musculature, and the sense organs. Fibres also interconnect the
two rings. The lower ring is the centre of rhythmic pulsations, i.e.,
it contains the pacemakers. Pulsation will continue in the bell as
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 long as any portion of the ring is intact. It is with the lower ring
that the statocysts are connected.

Sense organs:

The bell margin is richly supplied with sensory cells and also
contains two types of true sense organs, viz., light sensitive ocelli
and statocysts. The ocelli consist of patches of pigment and
photoreceptor cells organised either within a flat disc or a pit. The
ocelli are typically located on the side of the tentacular bulbs.

Statocysts are located between the tentacles or associated with

the tentacular bulb at the tentacle base. They may be either in the
form of pits or closed vesicles but in both cases, the walls contain
sensory cells with bristles projecting into the lumen. Attached to
the bristles are from a few to many calcareous concretions known
as statoliths.

The statocysts act as organs of equilibrium. When the bell tilts,

the statoliths respond to the pull of gravity and stimulate the
sensory bristles to which they are attached. The animal may then
respond by muscular contractions to bring itself back into a
horizontal position.

Reproductive organs:
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 The medusae are sexual or reproductive zooids possessing
gonads. The medusae are dioecious, they have either four testes
or four ovaries in the sub-umbrella just below the radial canals. A
gonad has an outer ectoderm and inner endoderm with
mesogloea between the two layers. The gonad has a small
diverticulum of the radial canal.

The germ cells of Obelia do not arise in the gonads, they arise
from interstitial cells of the ectoderm of the blastostyle where
they may be seen in various stages of maturation, then they
migrate into the medusa, then through the radial canals they take
up their position in the ectoderm of the gonads. When germ cells
mature, the gonads rupture and spermatozoa and ova are
discharged externally into water.

Life History of Obelia:

Fertilisation:

Fertilisation usually takes place in open sea water where the

gametes are set free. Sometimes, the sperms are carried into the
female medusae with water currents and there they fertilize the
eggs in situ. However, the parent medusae die soon after
liberating their respective gametes.

Development:

The zygote undergoes complete or holoblastic and equal

cleavage to form a single-layered blastula with a blastocoele.
Some cells migrate into blastocoele, eventually filling it
completely to form a solid gastrula known as stereo gastrula. Its
outer cell layer becomes the ectoderm and inner cell mass the
endoderm.
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 The gastrula elongates and. its outer layer of ectoderm cells
becomes ciliated, and now it is called planula. Soon, a cavity
called enteron develops in the solid endodermal cell mass by the
process of delamination and the planula becomes a two-layered
larva having an outer ciliated ectodermal cells and an inner layer
of endodermal cells.

The planula after a short free-swimming existence settles on

some solid object by its broader end. The free end forms a
manubrium with a mouth and a circlet of tentacles. Thus, a simple
polyp or hydrula is formed which grows a hydrorhiza from its
base, from which an Obelia colony is formed by budding.
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 The free swimming planula stage in the life history of Obelia,
helps in the dispersal of the species. The life history may be
represented as male and female gametes → zygote → planula
larva → hydrula → colony → sexual medusae → gametes → zygote
and so on.
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 Alternation of Generations and Metagenesis of
Obelia:

It is clearly evident from the life history of Obelia that there is an

alternation of polypoid and medusoid generations.

The polypoid generation is asexual and produces polyps and

blastostyles by asexual budding. The blastostyle also produces
medusae by asexual budding. The medusae do not produce
medusae but they give rise to gametes, which after fertilisation
develop into a polypoid colony from which medusae are produced
again by budding.

Thus, an asexual polypoid generation alternates with a sexual

medusoid generation. This phenomenon is known as alternation
of generations, till recently. The term alternation of generations
means that the individual exists in two distinct forms, which
alternate each other regularly in the life history.

One individual possesses the power to reproduce the other by

asexual reproduction, which again by sexual reproduction gives
rise to the next generation. Therefore, a true alternation of
generations is always between a diploid asexual and haploid
sexual generations, as is exhibited by fern plant.

But, in Obelia the condition is somewhat different and, therefore,

objections were raised to use the term alternation of generations
for it. Because, in Obelia, there are no true two generations to
alternate each other. The medusae are modified zooids capable of
free swimming existence and moreover they are not produced
directly from a zygote but are budded off from the blastostyle.

The gonads found in medusa are not formed in it but actually they
are formed in the ectoderm of blastostyle which later on migrate
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 into the medusa and become situated on its radial canals. Thus, it
is rather difficult to distinguish between sexual and asexual
generations. Hence the term metagenesis is used to replace the
term alternation of generations in Obelia.

Thus, in the life history of Obelia, there is a regular alternation

between fixed polypoid and free-swimming medusoid phases,
both of them being diploid.

Such an alternation of generations between two diploid phases is

known as metagenesis. Although, the phenomenon of
metagenesis is also reported in other groups of animals but it is
well represented by polymorphic hydrozoan like Obelia. Obelia
shows polymorphism in which the polyps are for feeding the
colony, blastostyles for budding and medusae for disseminating
gametes.

Advancement of Medusa over Polyp:

Medusa exhibits many features of advancement over polyp,

few of them are as follows:

1. The epidermis resembles the epithelium of higher Metazoa

forming a thin, protective and sensitive layer of small cells.

2. The enormous development of mesogloea reduces the gastro

vascular cavity or enteron to a system of canals and also provides
lightness which helps in buoyancy.

3. The nervous system shows differentiation into two nerve rings

constituting the central nervous system and nerve nets forming
the peripheral nervous system.

4. The marginal sense organs present at the bases of 8 tentacles

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 are of special advantage to the free swimming habit of the
medusa.

5. The mode of sexual reproduction provides wide dispersal of the

species due to its free swimming habit.

Similarities between Polyp and Medusa:

Striking as is the difference between polyp and medusa. They are

strictly homologous or typically similar structures. Both of them
are formed on the same pattern.

However, the features of similarity between them are listed

below:

1. Both are radially symmetrical.

2. Both are diploblastic with outer epidermis (ectodermal) and

inner gastro dermis (endodermal).

3. The mouth is homologous in both the cases; the mouth situated

on the hypostome in polyp is homologous with the mouth situated
on the manubrium of the medusa. Anus is absent in both the
cases.

4. The stomach, radial canals and circular canal of medusa are

homologous with the gastro-vascular cavity of the polyp. All
these are lined by gastro dermis and serve the purpose of
digestion and distribution of digested food.

5. Both are carnivorous; the food is captured and ingested with

the help of tentacles.

6. Digestion is extracellular as well as intracellular in both the

cases.
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 7. The outer, exumbrellar surface of the medusa is homologous
with the base of the polyp providing attachment with the parental
colony.

Derivation of Medusa from Polyp:

Striking as is the difference between polyp and a medusa, they

are strictly homologous structures, and the more complex
medusa is readily derived from the simpler polyp-form. The apex
of the umbrella of medusa corresponds with the base of a
hydranth. The mouth and manubrium are also homologous
structures.

Suppose the tentacular region of a polyp to be pulled out, as it

were, into a disc-like form and afterwards to be bent into the form
of saucer with the concavity distal, that is towards the
manubrium. The result of this to be a medusa-like body with a
double wall to the entire bell, the narrow space between the two
layers containing a prolongation of coelenteron and being lined
with gastro dermis.

From such a form the actual condition of things found in the

medusa would be produced by the continuous cavity in the bell
being for most part obliterated by the growing together of its
walls so as to form the endodermal lamella.

The cavity would remain only along four meridional areas, the
radial canals and as a circular area the circular canal close to the
edge of the bell. In this way a medusa is derived completely from
a polyp (Fig. 32.12).
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