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"PROTOZOA"
INTRODUCTION
The dual view of the living world, well manifested in the
work of Linnaeus ('Systema Naturae') with the proposal of 2
kingdoms, perdured almost until the early 70's in the last
century. If one organism was photosynthetic or was growing in
the ground, it was a 'plant' (Lt. 'planta' means a plant), and
it was moving freely, then it was an 'animal' (Lt. 'anima', that
has a soul: it feels and moves under its own impulse).
Despite

of

knowing

large

differences

between

the

'microrganisms'
in
terms
of
their
cellular
organization
(prokaryote vs. eukaryote), the proposed extension of the 2
classic kingdoms was accepted late in the scientific world.
It is not surprising, therefore, that this duality has
subordinated the study of the unicellular eukaryotes, either to
the
Botany
or
Zoology,
according
to
photosynthetic
or
heterotrophic attributes. Thus, in front of the Metaphytes or
multicellular plants, the Botany began with Protophytes or Algae
(including some forms of prokaryotes and Fungi). In a parallel
way, the Zoology separates the multicellular forms under the
name METAZOA, whereas the heterotrophic unicellular organisms
were grouped under a suggestive name: PROTOZOA.
The implications of a dual system, although convenient,
distorts the reality, because it is not offering a real
phylogenetic dichotomy. The change from prokaryote to eukaryote
has a larger significance for the living world than the change
from Plant to Animal. Moreover, the phylogenetic relationships
of unicellular eukaryotes are much more complex than previously
thought. In addition, there are some groups of eukaryotes, such
as flagellate euglenoids, with genera including species with
plant affinities, other species physiologically closer to

animals, and the rest they behave both like animals and plants
at the same time. For example, some dinoflagellates obtain 5% of
its energy requirements through photosynthesis, while the
remaining 95% follows heterotrophic metabolic pathways. Even
some flagellates can change from autotrophy to heterotrophy,
after maintaining in darkness for 24 hours.
All these contradictions encouraged the spread of the
proposal of Whittaker (1969), where the living world was divided
into two large super-kingdoms and 5 kingdoms (see fig. 1).
PROTISTA was placed at the base of the three large kingdoms of
multicellular eukaryotes: PLANTAE, ANIMALIA and FUNGI. As
consequence, all unicellular eukaryotes contain the groups
classified
as
PROTOPHYTA,
PROMYCOTA
and
PROTOZOA.
The
heterogeneity of some taxa of PROTISTA has influenced new
proposals of kingdoms of the living world. As we see, the
validity of the term 'protozoan' is not greater than that
achieved by the term 'worm', and yet it is the subject of this
chapter deal with protozoans. A recent classification system
proposed for PROTISTA recognized up to 27 phyla, more than half
exhibit autotrophic metabolic pathways, and the remaining 13 are
heterotrophs. Of these 13 heterotrophs, 9 are endoparasites or
endosymbiotic.
Three
phyla
(DINOPHYTA,
EUGLENOPHYTA
and
CRYPTOPHYTA) include species
botanists and zoologists.

which

have

been

disputed

by

Most of these protozoan protists are phylogenetically very


distant from metazoans. In fact, they do not share positive
characteristics
with
metazoans.
By
using
some
negative
functional characteristics like heterotrophy, we would be forced
to relate the
heterotrophic.
The
kingdoms

METAZOA

with

PROMYCOTA,

fundamental difference
of eukaryotes is the

since

they

are

also

between protists and other


character of unicellularity

(solitary or colonial) vs. multicellularity. The protist cell is


able to perform all the functions, necessary for its life

survival
and
reproduction.
By
contrast,
multicellular
eukaryotes, carry out all these general functions by cell groups
integrated in different tissues, even in complex organs.

Fig. 1. The system of 5 kingdom and its relationship with the


duality plant-animal

As we can see this difference is only semantic, since all


the animal cells have a common origin in one unique diploid cell
(zygote), and just at that time the zygote is so totipotent as a
protist cell.

Fig. 2. Main evolutionary lines to explain the multicelluarity


origin from unicellular forms.

There are theoretically three ways in which a multicellular


animal could evolve from unicellular ancestors. First, there are
authors who consider the symbiosis as a mechanism for
multicellularity from different groups of protozoans (Fig. 2a).
Secondly, other authors consider the colonialism as a way to

obtain an incipient multicellularity, since by simple mitosis we


can obtain a colonial form, in the case the cells are not
finally separated (Fig. 2b). Thirdly, there are followers who
consider the cellularization as a useful mechanism to obtain
multicellular forms from a multinucleate protozoan, that
secondarily could promote cells by internal membrane partitions
around each of its nuclei, becoming internally compartmented
(Fig. 2c).
The first of these three potential mechanisms presents
serious genetic problems, since has to overcome differences
within the protozoans to integrate into a single multicellular
organism. In the case of lichens (algae + fungus), each organism
reproduces separately to end his life in a symbiotic form.
Regarding the third mechanism, there is no evidence that occurs
within the present o past protozoans. Consequently, it seems an
evolutionary path not very suggestive. However, in relation
the second mechanism, we find many protists able to
colonies,
and
to
promote
a
basic
and
primitive
differentiation. Therefore, colonialism represents a

with
form
cell
more

plausible mechanism to explain the origin of multicellular forms


from unicellular ancestors. In fact, it is difficult to
distinguish
between
a
colonial
protozoan
and
a
simple
multicellular organism. In the last one, we can observe there is
a fragile coordination of their cells. Moreover, the cell
differentiation is not only confined to multicellularity, since
there are colonial protozoans with several cell types. Actually,
it is more a semantic problem, if we consider there is a
continuous line from unicellularity to multicellularity thanks
to colonialism. Of the 27 phyla of protists recognized in the
latest classifications, 16 phyla have colonial species, and at
least 3 phyla show a colonial organization which reaches the
multicellularity in a strict sense.
In conclusion, the 'protozoan' grouping is today obsolete,
since
it
represents
a
heterogeneous
association
(not
phylogenetic) of different heterotrophic protists. We should be

aware that protists have had a complex evolutionary history on


its
own.
Of
these,
only
one
group,
CRASPEDOPHYTA
(CHOANOFLAGELLATA), shares several characters with METAZOA. This
fact has allowed to use choanoflagellates as an appropriate
model to explain the origin of animals. Here, we will discuss
the protozoans as a short introduction to metazoans.

DIAGNOSIS
Protozoans are eukaryotic, unicellular, with 1 to several
nuclei, capable of forming colonies. Most protozoans are
microscopic. In the colonies of protozoans there is an incipient
cell differentiation in germ cells and somatic cells (first sign
of a cellular labour division). However, protozoan cells are
unable to form tissues.

GENERAL CHARACTERS
From a zoological point of view, it should be noted that a
large number of protozoan species are colonial and some species
exhibit clearly multicellular stages during their life cycle.
The reason for further consideration of these organisms as
protozoans is given: (i) primarily by an easy recognition of
their relationship with other unicellular forms, (ii) by lacking
somatic cell differentiation, and finally (iii) by lacking
embryonic development. Metazoans are multicellular by a somatic
cellular
differentiation,
which
appears
well
defined
by
embryonic processes. The difficulty in establishing a neat
border between colonial protozoans and metazoans should be
solved conceptually with the following point of view: the
singularity of the metazoans is not their multicellularity
(which is shared with groups of protozoans), but better by the
evolutionary significance the metazoans are going to give to the
multicellular character along evolution.

Briefly,

we

can

list

the

singular

characteristics

protozoans when compared with metazoans:


1. Unicellular and totipotent; some are colonial
ability to undergo neat multicellular stages.
2. Lacking blastodermic layers; there are no

with

of
the

embryonic

processes.
3. Lacking tissues or organs, although they exhibit very complex
cell organules.

Fig. 3. Major types of PROTOZOA: a)


(amoebae); c) ciliates and d) sporozoans.

flagellates;

b)

rhizopods

CLASSIFICATION
Protozoans can be grouped under four basic forms (see fig.
3). This classification is useful in a simplified study of the
protozoans from the external point of view of metazoans. The
groups are: (1) Amoebae or rhizopods: its main characteristic is
the possession of small cell appendages called pseudopods. (2)
Flagellates: whose main characteristic is the possession of
flagella. (3) Ciliates being its main characteristic the
presence of numerous cilia, that occur in high rows along their
bodies. Each cell contains nuclei of two types: a macronucleus
and a micronucleus. (4) Sporozoans, endoparasites, and its main
feature is the use of spores for transmission of infections.
Often, groups 1 and 2 are further grouped under the phylum
Sarcomastigophora, the group 3 within the phylum Ciliophora, and
group 4 of the "Sporozoa" has been revealed as extraordinarily

complex from an ultrastructural point of view, proposing up to 4


phyla: APICOMPLEXA, MICROSPORA, ASCETOSPORA and MYXOZOA. We are
only dealing with the first phylum because its importance to
explain the origin of METAZOA.

Phylum SARCOMASTIGOPHORA (FLAGELLATA + RHIZOPODA)


(Approx. 25.000 sp.)
(gr. sarx: meat; mastix: whip; pherein: carry)
This phylum comprises protozoan species with pseudopodia or
flagella for trophic / locomotion purposes. It includes the
amoebae and flagellates from the old systems of classification.
The reason for its grouping is based on the fact that many
flagellates are able to form pseudopods and some amoebae under
favourable environmental conditions are capable of using
flagella (Fig. 4).
Within the sarcomastigophorans, we consider only the
subphylum Mastigophora (Flagellata) by its importance to explain
the origin of METAZOA.

Subphylum MASTIGOPHORA (= FLAGELLATA).


- Their representatives exhibit from 1 to several flagella.
- Usually have a single nucleus.
- According to the nutrition preferences, they are two major
classes:
- Class PHYTOMASTIGOPHOREA (with chloroplastes)
- Class ZOOMASTIGOPHOREA (without chloroplastes)

Fig 4. Naegleria. A): flagellate forms; B) the ameboid form can


change to a flagellate form and develop into a cyst finally.

The first class, as its name indicates, shows affinities


with algae within the Botany. The class has up to 20 orders that
are differentiated by their colour and placement of the
flagella. Examples of this class are dinoflagellates, Euglena
and volvocids. The last ones are particularly interesting as an
illustrative model to explain colonialism and multicellularity.

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Fig. 5. Colonial forms within the Volvocida. Starting from one


single cell of Chlamydomonas (1) there are colonies of growing
individuals like in Gonium (2) and Pandorina (3). Eudorina (4)
shows somatic cells (in black) and Volvox (5) shows many somatic
cells (in black) together with macrogametes and microgametes.
Colonies of Volvox aureus (freshawater species) may measure from
200 1000 microns, and are integrated by approx. 500 cells, with
a central gelatinous mass. This sphaerula swims thanks to a
coordinated movement of every cell with two flagella.

In fig. 5 it is illustrated the formation of colonies


within the VOLVOCIDA. Nevertheless, there are difficulties in
explaining the derivation of the animals from this model, mainly
for the possession of a cell wall and chromatophores.
The class of zoomastigophorans has no species capable of
photosynthesis. Mostly are parasites.
However, within the class zoomastigophorans, there is an
order with free living representatives, choanoflagellates, which
serve as a suggestive model for the origin of METAZOA. They have
a cell with a single flagellum in the apical part surrounded by
a collar of fine pseudopods. The function of the flagellum is to
create water currents, which have to pass through a mesh created
by the pseudopods. It retains the food particles suspended in
the water column (microfiltration). Choanoflagellates are widely

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distributed, with both solitary and colonial forms, as well as


sessile and motile. Marine species often secrete silica
spicules, which can form a protective capsule called lorica
(Fig. 6). The ultrastructure of the apical collar and flagellum
of choanoflagellates is similar to the sponge choanocyte cells
(Fig. 7). Since both groups are capable of secrete silica
spicules, it is not surprising the proposal of choanoflagellates
as precursor forms of the sponges. Especially, colonial
choanoflagellates like Proterospongia (Fig. 6) and Sphaeroeca,
and the sessile form Codosiga, have been used as intermediate
forms between protozoans and sponges.

Fig. 6. a) Salpingoeca amphoroideum is a sessile marine form


surrounded by a lorica of silica spicules. In its apical part
there is collar with 25 pseudopods. b) Proterospongia haeckeli
shows both flattened and sphaerical colonies, and inhabits
freshwater. Their colonies can be formed by up to 100 cells
which remain trapped in a central gelatinous matrix. Some
tendencies towards the cell differentiation are shown in fig. d,
where
are
amoeboid
cells.
c)
Acanthoecopis
apoda is a
choanoflagellate with silica spicules forming the exoskeleton.
d) Development of Proterospongia haeckelii. It starts from a
single sessile cell without collar. Later on, it forms a colony
with an incipient cell differentiation.

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Fig. 7. Similarity
protist cell.

of

the

choanocyte

and

choanoflagellate

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