Metazoa
Metazoa
Metazoa
ANUPAM GHOSH
The animal kingdom comprises of a vast and heterogenous assemblage of animals. The
range in size, in structural diversity and in adaptations to different modes of existence is
enormous. However, in spite of such diversity the animal kingdom shows certain
distinctions of categorization on the basis of their origin and embryological
development. The first step in this categorization is the broad divisions of the animal
kingdom into single celled animals or protozoa and multicellular animals or metazoan.
Evolution of the metazoans:
The sequence of events in the metazoan evolution began at the protozoan level and can be sequenced as
follows:
1.The first step in the evolution of metazoans involved the development of an initial loose organization and
partial specialization of cell types (particularly in reproduction) as can be seen in case of mesozoans and
placozoans.
2.At the next step came a stage where by specific body functions were restricted to specific cells, which was
restricted in sponges.
3.Next came the stage of advanced cell specialization to produce true tissues. This stage highlights a
sequence of events that involved not only development of tissues but also development of embryonic germ
layers. The animals of this stage have two germ layers and radial symmetry. Example includes ctenophores
and cnidarians.
4.In the next step, evolves pseudocoelomates characterized by early cell cleavage. They lack viable primary
larval phases and embryonic development is poorly developed. Example of this stage includes nematodes,
Acanthocephala, Rotifera and Kinorhyncha.
5.Later on, developed the acoelomates and coelomates. They have a highly regulated and determine mode of
development as evidenced by spiral cleavage and mesoderm formation. They are typically characterized by
possession of distinct larval stages. Three groups resulted from this stage-
a)the acoelomates who never developed complex body plans due to lack of body cavity, and
b)the other two lineages were based on a coelomic body plant- protostomia and deuterostomia
Theories regarding the origin of Metazoa:
Although there are a number of theories regarding the origin of metazoan, but
most zoologists are of the opinion that metazoan has originated from single
celled protozoa. Such an opinion has been put forward by several scientists,
such as, Haeckel (1840), Metschnikoff (1887), Hymann (1940), Hadzi (1953),
Hanson (1977) etc. The most modern theories which have been put forward in
support of single celled origin of Metazoa are – Syncytial theory, Poly phylectic
theory and colonial flagellate theory.
a) Syncytial Theory: The theory was proposed by Hanson (1958) and Hazda
(1963) and proposed that multinucleate ciliates must have been the ancestors
of metazoans. Ciliates possess high grade of organelle development and body
organization among protozoans. By partitioning the multinucleated cytoplasm
with cell membranes the ciliate body could have become multicellular and gave
rise to metazoa.
b) Polyphylectic theory: Metazoans are polyphyletic, derived from more than
one group of unicellular organisms. Sponges, cnidarians, ctenophores and
remaining eumetazoans each evolved independently.
c) Colonial flagellate theory: Historically, Haeckel’s Gastraea theory was the first widely accepted
model of the origin of animals. According to Ernst Haeckel, the transition from unicellularity to
multicellularity proceeded through two consecutive stages (Fig. 1A). First, unicellular flagellates
aggregated to form a hollow ballshaped floating colony of identical cells. This hypothetical stage was
named the Blastaea to reflect its similarity with the blastula, an early stage in animal embryogenesis.
Outside the Metazoa, ball-shaped colonies of flagellated cells are observed in the green algae Volvox
and the choanoflagellate Sphaeroeca volvox, to name a few. Ultrastructural evidences points to the
choanoflagellates as the best candidates, as they have mitochondria and ciliary rootlets that are very
similar to those in metazoan cells.At the second stage, the homogenous cellular wall of the Blastaea
invaginated to form a second layer, the precursor of the primary gut. After this important event the
Blastaea acquired primary cell differentiation. Haeckel named this hypothetical ancestral form the
Gastraea, again to indicate its similarity with the gastrula stage in animal embryogenesis. According
to Haeckel, extant cnidarians and sponges are the first direct descendants of the ancestral Gastraea,
because their body plans derive from two embryonic layers. The presence of a uniform blastula and
gastrula with differentiated ectoderm and endoderm were postulated by Haeckel to be the
characteristic of all Metazoa.