Kingdom Animalia

PHYLUM ANNELIDA

INTRODUCTION

Of the three major phyla of protostomes, only the arthropods and annelids exhibit
metamerism, the division of the body into segments. Segmentation is advantageous during
development, where greater efficiency is obtained by constructing a whole organism out of
identical modules. In the adult, locomotor activity is enhanced because of the independent
nature of each segment and the flexibility afforded by a series of segmented pans.
Segmentation also gives these phyla a survival advantage. Since many segments are similar
to other segments in form and function, damage to one or several segments does not
necessarily compromise body functions.

The phylum Annelida (segmented worms) is diverse, containing the earthworms, leeches,
and marine worms. They are all triploblastic, bilaterally symmetrical, and eucoelomate, In
addition, annelids exhibit a body wall with both longitudinal and circular muscle layers
(which, along with segmentation mentioned above, allows these animals to be quite
mobile), a complete digestive tract, a nervous system showing some degree of
cephalization, a closed circulatory system, and an excretory system (based on nephridia).

There are three major classes within the phylum Annelida, described below.

Class Polychaeta - mostly marine worms, such as Nereis (the clamworm)

Class Hirudinea - the leeches (predominantly freshwater), such as Hirudo
Class Oligochaeta - mostly freshwater and terrestrial worms, such as
Lumbricus (the earthworms)
 PROCEDURE:

In this exercise, we will examine the external and internal structures of

Lumbricus. Nereis and Hirudo are available as demonstrations.

EXTERNAL ANATOMY

Obtain an earthworm from the supply and place the animal in a dissecting
tray. You may need a dissecting scope to fully appreciate the external
anatomy. The most obvious external feature is the clitellum, a swollen area in
the anterior third of the specimen. This region functions in reproduction by
secreting a mucous which holds the participants together during sperm
exchange.

Orient the worm dorso-ventrally by locating the setae. Run your fingers along
the animal to feel the rough texture produced by the setae. Four of these
structures are found on the ventral surface of each metamere. They provide
traction during locomotion.

Using the clitellum as a landmark, identify the anterior and posterior ends of
the worm. The anterior end has a mouth opening, covered by a lobe termed the
prostomium. The posterior segment bears the anus.

Starting with the segment that holds the mouth, locate segment 14. Observe
the openings for the oviducts on the ventral surface. Find the sperm ducts on the
ventral surface of metamere 15.

INTERNAL ANATOMY:

Pin your worn , dorsal side up, near one edge of a dissecting pan. Using fine
scissors or a sharp scalpel, make a shallow longitudinal cut along the dorsal
surface of the worm. (Be sure not to cut into the dorsal vessel and the
intestinal tract, which are located just below the body wall.) Note that the
body wall is held in place by septa (internal divisions between metameres). Cut
the septa along the length of the worm on both sides of the intestine. These
septa divide the coelom into separate cavities. Pin the body wall to the bottom
of the pan, inclining the pins away from the worm. Keep your preparation
moist at all times.

Digestive system:

Observe the digestive tract running from mouth to anus. Anteriorly there are a
number of modifications of the tract. Just posterior to the mouth opening is
 the muscular pharynx. The torn muscles associated with the pharynx were
attached to the body wall. When these muscles contract, food particles are
sucked into the mouth. The esophagus is posterior to the pharynx and is
surrounded by cream-colored bodies that will be studied later. The esophagus
expands into a thin-walled storage structure, the crop. Probe the wall of the
crop gently and note its texture. Just posterior to the crop is the muscular
gizzard. This is a grinding structure with thick, muscular walls (obvious by
gentle probing). Food is passed from the gizzard to the intestine, where further
digestion and absorption occur. The intestine ends at the anus.

Reproductive system:

Earthworms are monoecious, with cross-fertilization the rule. The gonads

(testes and ovaries) may be too small to see, but many of the associated
reproductive structures are clearly visible. Note the large, cream-colored
structures associated with segments 9-12. These are seminal vesicles. Testes
are associated with seminal vesicles. Sperm are passed from the testes to the
seminal vesicles for storage prior to copulation. During copulation sperm exit
through a duct system opening at segment 15. The ovaries are located in
segment 13. Their duct system opens to the outside in segment 14. Two pairs
of small, round, cream-colored structures are present on the ventro-lateral
body wall in segments 9 and 10. These seminal receptacles receive sperm during
copulation.

During copulation, two worms line up facing in opposite directions, segments 9

and 10 of each opposite the clitellum of the other. The anterior halves of the
worms are wrapped in a mucous sheath. Sperm from the seminal vesicle of
each passes into the seminal receptacle of the partner. When worms separate,
sperm transfer has been mutual. Each worm then forms a second mucous
sheath at the clitellum. Eggs are discharged into the sheath and the worm
begins backing out of the sheath. On reaching segments 9 and 10, sperm from
the seminal receptacles are discharged into the sheath and fertilization takes
place. The sheath is closed and shed as it reaches the head, forming a cocoon
within which direct development (without a larval stage) occurs.

Excretory system:

Nephridia are organs of excretion in the annelids. Flood your worm with water.
Observe the body wall under a dissecting microscope. The nephridia are coiled
tubules with an expanded funnel-shaped nephrostome. The nephrostome is
attached to the septum dividing two segments and opens into the anterior
segment. The tubule opens to the outside through the body wall within the
posterior segment. Filtration of the blood across the tubule wall can occur
because of the close association between capillaries and the nephridium.

Circulatory system:
 Circulation in the earthworm is through a series of closed vessels. The two
main vessels that can be seen in your dissection are the dorsal and ventral blood
vessels. These vessels are the main pumping structures. In the dorsal vessel,
blood moves anteriorly. The dorsal vessel is the dark line running along the
dorsal surface of the digestive tract. In the posterior third of your worm,
carefully cut through and remove about three centimeters of the digestive
tract. The ventral blood vessel can usually be seen adhering to the segment of
intestine removed. In the ventral vessel, blood moves posteriorly. Segmental
branches off the ventral vessel supply the intestine and body wall with blood.
These branches eventually break into capillary beds to pick up or release
nutrients and, oxygen. Gas exchange occurs between the capillary beds of the
body surface and the environment. Oxygen is carried by the respiratory
pigment hemoglobin, which is dissolved in the fluid portion of the blood. From
these capillary beds, blood is collected into larger vessels that eventually unite
with the dorsal vessel. At the level of the esophagus, segmental branches are
expanded into five pairs of aortic arches, or what have been called "hearts".
They are dark, expanded structures on either side of the esophagus. Although
these are contractile, they only function in pumping blood from the dorsal to
the ventral vessels.

Nervous system:

In the region of your worm where the intestine was removed, locate the ventral
nerve cord with its segmental ganglia (seen as slight swellings). Nerve fibers
from segmental ganglia innervate structures in each segment. Extend your
mid-dorsal incision to the prostomium and locate the paired supra-esophageal
ganglia anterior and dorsal to the pharynx. Circum-pharyngeal connectives lead
from the supra-esophageal ganglia to the ventral nerve cord.

CROSS-SECTION:

Using the lowest power of a compound microscope, examine a cross-section of

Lumbricus. Note the noncellular, protective cuticle covering the epidermis. Just
below the epidermis, note the circular muscle and, below that, the longitudinal
muscle. The mesodermal lining of the inner body wall is the peritoneum.
Covering the dorsal ve
 ssel and the intestine are cells called chlorogogue cells. These are involved with
glycogen and fat synthesis and urea formation. Folding to the interior of the
intestine is the typhlosole, which increases surface area for secretion and
absorption. Two lateral neural blood vessels and one subneural blood vessel are
associated with the ventral nerve cord. Depending on the section studied you
may also see nephridia and setae.
Earthworms

The subclass Oligochaeta contains all the animals commonly thought of as 'earthworms'. Long
thin worms with no obvious appendages to their bodies and greatly reduced heads so that when
the animal is still it is not sometimes obvious which end is the head and which is the tail. As a
group they are all morphologically similar and taxonomic division is often made far more on the
basis of internal characteristics, particularly the positioning of the genitalia.

The Oligochaeta are the second largest group of the Annelida, with 3,100 known
species they make up about one third of the phylum. within this diversity of species
there are aquatic forms, both freshwater and marine and also many terrestrial
species. In terms of diet, the smaller species are often predatory while the larger
species are soil or mud feeders. There are also a few parasitic species.

The 'Oligo' in Oligochaeta means 'few' just as the 'Poly' in Polychaeta means many,
thus the Oligochaeta are the animals with few chaetae, or few bristles. Normally, as
in the common earthworms, Oligochaetes have 8 small chaetae per body segment.
These are usually arranged in four groups of two around the body. The Oligochaetes
are also distinguished from the Polychaeta in that they are hermaphroditic and in
that they possess a clitellum as adults. This is an organ situated on the anterior
section of the animal that is important in reproduction.

What the Oligochaeta lack in terms of species number, in comparison with the
Polychaeta, they make up for in numbers of individuals. In some places in Britain
there are 1 ton common earthworms per acre which move well over 10 tons of soil
per year. Common earthworms are quite large, in comparison with many soil
animals and the total numbers present in a given soil vary enormously in different
soil types, habitats and soil usage regimes. Generally speaking deciduous woodland
and orchard soils support the highest numbers, also temperate soils tend to support
higher number than tropical soils. Over all soil types the range per square metre of
soil is from less than 1 to more than 800 individuals. Other groups of Oligochaetes
are smaller and can be packed more densely into a similar area, thus in the mud of
the River Thames you can find more than 40,000 Oligochaetes per metre square,
most of which will be Tubificids, while further inland it is not difficult to find up to
300,000 or more Enchytreids per metre square of moorland soil.
Earthworms are enormously important in the construction and fertility maintenance
of the soil and were described by Aristotle as 'the intestines of the earth'. Numerous
soil scientists have been equally fascinated by the amount of work done by them,
Charles Darwin said of them "It may be doubted whether there many other animals
which have played so important a part in the history of the world as these lowly
organised creatures".

Earthworms are miniature topsoil factories, earthworms make soil, all other living
things eventually pass through an earthworm on the way to becoming soil, and it is
likely that nearly every atom in your body has been in an earthworm's stomach
before it was part of you, (some salts and ions, and some water are the exceptions).
Earthworm castings are rich in all the minerals necessary for plant growth in a water
soluble form so that they are immediately available for plant use. All the soil you
have ever seen, at least in temperate climes, has passed through the stomachs of
numerous earthworms to become what it is. The best way to deal with those
unwanted earthworm casts on your lawn is to collect them and use them as potting
compost, no manufacturer can make anything better suited to the growth of
seedlings.

It has been scientifically proven that earthworms increase the productivity of many
soils, in some cases doubling or tripling crop yields. They do this by improving the
structure of the soil, by bring nutrients up to the surface layers of the soil from
deeper down and assisting in the break down of organic matter in, or on the surface
of the soil.

This has resulted in a huge market in worms springing up in the second half of the
20th century in Western Europe and USA. In many cases this has been good, but in
others, ignorance has resulted in the efforts and money spent being wasted. Soils
have a definite worm carrying capacity which relates directly to the amount of
organic matter in, and regularly added to, them (yearly in natural environments).
Adding more worms than the soil can carry, or adding worms to soils poor or
deficient in organic matter will result in only a small temporary increase in fertility
resulting from the nutrients released from the dead worms and not from the work
they have done. Further more not all worms are suitable for all soils and not all
worms are active in soil creation, in particular Eisenia foetida, a commonly sold
species which is very useful in composts and dung recycling is useless in, and will
not survive in, ordinary field soils.

Follow this link to learn about worm farming for kids.

Nervous System

The nervous system consists of a brain, which is found in the prostomium and
connected to the ventral nerve cord, and the sub-pharyngeal ganglia, by a special
set of nerves called the circum-pharyngeal connectives. The ventral nerve cord is
surrounded by a fibrous sheath. The nerve cord contains two sorts of nerve fibres;
normal nerve and giant nerves. The giant nerves are only important during rapid
escape manoeuvres when the animal needs to react very quickly. The ventral nerve
cord runs the whole length of the animal and gives rise to several sets of nerves in
each segment. In the more active forms each segment has its own small ganglia,
but in the more sedentary forms these are absent. Also the more active the lifestyle
of an animal the larger its brain, because of the need to interpret a much greater
input of sensory information as well as the need to co-ordinate more varied and
complicated movements.

Blood and Circulation

The blood system is a closed system, meaning that they have blood vessels through
which the blood flows. There are two main longitudinal blood vessels, a dorsal one
and a ventral one, as well as three smaller longitudinal vessels, two lateral neural
vessels and one sub-neural vessel. These vessels have circular muscles around
them which can contract rhythmically to keep the blood moving around the body.
The vessels also contain valves which ensure the blood only flows in one direction.
Blood flows from the head to the tail in the ventral vessel and back, from the tail to
the head, in the dorsal vessel. In each segment a number of smaller lateral (side)
vessels branch off from the main vessels to supply the sections of that segment.
The blood of annelids contains haemoglobin, the same respiratory pigment as in
humans, it is this that makes their blood the same red colour as ours.

Gaseous Exchange

Gaseous exchange normally occurs over the whole of the animal's body. Most
aquatic oligochaetes, being smaller than polychaetes, have no need of special
additional respiratory organs. Becausegaseouss exchange is far greater in the
presence of moisture terrestrial species, secrete moisture in the form of coelomic
fluid from the dorsal pores, mucous from the epidermal mucous glands and the
excretions of the nephridia.

Excretion and Osmoregulation

Excretion of metabolic wastes is through the action of nephridia. These are long
coiled tubes which have many cilia lining their internal surface. In Lumbricus these
are longer and more complicated than in the polychaetes and the tube is divided
into three distinct parts, the innermost section of which is far wider than
normalnephridiall tubes. Both blood and coelomic fluid enter these nephridia where
nutrients, water and salts are removed before the remaining wastes are passed out
through the nephridiopore. Most body segments, except the first three and last one,
have their own pair of nephridia. Nephridia also serve as organs of osmoregulation
and in species that live in fresh water, or environments with variable salinity, the
nephridial tubes are longer to help them deal with the greater osmotic potential
occurring between their inner body fluids and the fluids of the environment they are
living in.
Reproduction in Earthworms

Oligochaetes are hermaphrodites, with separate testis and ovaries. The sexual
organs, and the ducts that lead to and from them are situated in the anterior (front)
part of the animal, normally between segments 7 and 15. The actual placement of
the reproductive organs, including the openings of the ducts, which are normally on
the same segments are important in classification. In Lumbricuscus terrestris, a
common worm in Western Europe, the testis are in segments 10 and 11, the
seminal vesicles in segments 9,11, and 12, while the vas deferens opens on
segment 15. The ovaries are found in segments 13 and the oviduct opens on
segment 14. The sexual organs and their ducts are paired, one on each side of the
worms body. Sperm travels from the opening of the vas deferens to the clitellum,
segments 32-36, along two seminal grooves.

Copulation occurs on warm damp nights. The worms lie head to tail and side by
side. In this way the clitellum segments of each animal are opposite the segments
containing the sexual organs of the other. The clitella secrete a mucous tube that
surrounds the worm from before the first reproductive segment to the clitella
segments (segments 8 to 36 in L. terrestris). Sperm received from the partner worm
is stored in the spermathecal openings (segment 9 in L. terrestris) and then the two
worms separate.

Each worm now secretes a new mucous tube, one that is enriched with albumin
from the clitellum and wrapped in membranene. The eggs (5 - 16 in L. terrestris)
are shed into this tube along with some sperm. the worm then backs out of the tube
which now becomes an egg cocoon. Fertilisation occurs inside the cocoon. The
cocoon is left under the ground, or attached to plants under the water and often
changes shape, becoming darker, smaller and harder. In L. terrestris only one egg
survives the juvenile stage to emerge from the cocoon, but in other species more
than one will survive. The exact detail vary from species to species but the general
pattern remains the same.