CHAPTER 3 – UNICELLULAR EUKARYOTES o The presence of at least one motile stage

in the life-cycle
Part 1: Overview
Most ingest their food
A unicellular eukaryote is a complete organism The term “Protozoa” is not accurate
in which all life’s activities occur within the limits
ECOLOGICAL SIGNIFICANCE
of a single plasma membrane.
Unicellular eukaryotes are found wherever life
exists. They are highly adaptable and easily
moved from place to place.
They require moisture, whether they live in
marine or freshwater habitats, soil, decaying
organic matter, or plants and animals. They may
be sessile or free swimming, and they form a
large part of the suspended plankton. Parasitic forms cause some of the most
The same species are often found widely important diseases of humans and domestic
separated in time as well as in space. Some forms animals.
span geological eras exceeding 100 million years.
FEEDING METHODS

The two concepts, plantlike and animal-like,

refer in part to the way that food is gathered.
Plants are typically autotrophic, meaning that
they synthesize their own organic constituents
from inorganic substrates. Photosynthesis is one
kind of autotrophy.
Animals are typically heterotrophic, meaning
that they obtain organic molecules synthesized
by other organisms.
Heterotrophic unicellular eukaryotes may ingest
their food in a soluble form or in a particulate
form.
PHAGOCYTOSIS – way to ingest particulate form
of food.
o infolding or invagination of the plasma
membrane to surround a visible food
particle.
PHAGOTROPHS (holozoic feeders) – feed on
visible particles
OSMOTROPHS (saprozoic feeders) – feed on
soluble food

PROTOZOA

Not yet animals

The assemblage of unicellular eukaryotic
organisms was called Protozoa
The inclusion of “zoa” in the name referred to
two animal-like features:
o Absence of cell wall
 A flagellum propels water parallel to the main
axis of the flagellum.

Beating of flagella either pushes or pulls

the organism as seen in the Euglena
through its medium
Cilia propels the organism by a rowing
mechanism

LOCOMOTION METHODS

Originally, the means of locomotion was used to

distinguish unicellular eukaryotes.

Pseudopodia are extensions of the cell

cytoplasm used in locomotion and feeding.
Ectoplasm appears more transparent (hyaline)
by light microscopy, and it bears the bases of the
cilia or flagella.
Ectoplasm is often more rigid and is in the gel
state of a colloid, whereas the more fluid
endoplasm is in the sol state.
Pseudopodia vary in composition and are of
several types.

Some investigators have preferred to call them

both undulipodia (L. dim. of unda, a wave, + Gr.
podos, a foot).
A cilium propels water parallel to the surface to
which the cilium is attached.
 o Reticulopodia are distinguished from
filopodia in that reticulopodia
repeatedly rejoin to form a netlike
o The most familiar are lobopodia, which
mesh, although some biologists consider
are rather large, blunt extensions of the
the distinction between filopodia and
cell body containing both endoplasm
reticulopodia artificial.
and ectoplasm

o Axopodia are long, thin pseudopodia

o Filopodia are thin extensions, usually supported by axial rods of microtubules
branching, and containing only
ectoplasm. They occur in some amebas,
such as Euglypha
 described as compact or condensed because the
chromatin material is more finely dispersed and
clear areas cannot be observed with the light
microscope.

MITOCHONDRIA

A mitochondrion is an organelle used in

recovering energy from carbon bonds of fuel
molecules where oxygen serves as the terminal
electron acceptor.
It contains DNA.
Cristae, the internal membranes of a
mitochondrion are of variable form, being flat,
tubular, discoid, or branched (ramifying). The
form of cristae is considered a homologous
character (inherited from a common ancestor)
and, in conjunction with other morphological
features, is used to describe particular taxa.
Part 2: Functional Components of Unicellular
Eukaryotes

NUCLEUS

As in other eukaryotes, the nucleus is a

membrane-bound structure whose interior
communicates with the cytoplasm by small
pores.
Within the nucleus the genetic material (DNA) is
borne on chromosomes. Except during cell
division, chromosomes are not usually
condensed in a form that can be distinguished,
although during fixation of the cells for light
microscopy, chromosomal material (chromatin)
often clumps together irregularly, leaving some
areas within the nucleus relatively clear
(VESICULAR APPEARANCE).
Within the nucleus, one or more nucleoli
representing active transcription of ribosomal
RNA are often present.

o Hydrogenosomes are organelles that

Only genes of the macronucleus
transcribed. Macronuclei of ciliates
perform a respiratory function in the
absence of oxygen and are assumed to
have evolved from mitochondria.
o Hydrogenosomes and other
are mitochondrial derivatives that lack DNA
are are collectively called mitosomes.
 o Kinetoplasts are also assumed to be
mitochondrial derivatives. They contain
masses of circular DNA molecules and work
in association with a kinetosome, an
organelle at the base of a flagellum.
PHAGOCYTOSIS
GOLGI BODY, PLASTIDS, AND EXTRUSOMES
Golgi Complex
o The Golgi complex is part of the
endomembrane system that participates in
cellular secretory processes and intracellular
digestion of ingested food vacuoles.
o Golgi complexes are also called
dictyosomes.
o Parabasal bodies are similar structures with
potentially similar functions
Plastids
o Plastids are organelles containing a variety
of photosynthetic pigments.
o The original addition of a plastid to
eukaryotic cells likely occurred when a
cyanobacterium was engulfed and not
digested (secondary endosymbiosis).
o Chloroplasts contain different versions of
chlorophylls (a, b, or c), but other kinds of
plastids contain other pigments.
o For example, red algal plastids contain
phycobilins. Particular pigments shared
among unicellular eukaryotes may indicate
shared ancestry, but plastids also could have
been gained by secondary endosymbiosis
(one organism lives inside the other through
engulfment).
Extrusomes
o This general term refers to membrane-
bound organelles in unicellular eukaryotes
that are used to extrude something from the
cell.
o The wide variety of structures extruded
suggests that not all extrusomes are
homologous.
o The ciliate trichocyst is an extrusome.
In holozoic nutrition, food particles are brought
into a cell by phagocytosis where extension or
invagination of the plasma membrane
surrounds a food particle.
As the invagination extends farther into the cell,
it is pinched off at the surface. The food particle
thus is contained in an intracellular, membrane-
bound vesicle, a food vacuole or phagosome.
Lysosomes, small vesicles containing digestive
enzymes, fuse with the phagosome and pour
their contents into it, where digestion begins.
As digested products are absorbed across the
vacuolar membrane, the phagosome becomes
smaller. Any undigestible material may be
released to the outside by exocytosis, the
vacuole again fusing with the cell-surface
membrane.
In most ciliates, many flagellates, and many
apicomplexans, the site of phagocytosis is a
definite mouth structure, the cytostome.
In amebas, phagocytosis can occur at almost any
point by envelopment of a particle with
pseudopodia.
Many ciliates have a characteristic structure for
expulsion of waste matter, the cytopyge or
cytoproct, found in a characteristic location. In
some, the cytopyge also serves as the site for
expulsion of the contents of the contractile
vacuole
REPRODUCTION IN UNICELLULAR EUKARYOTES

Sexual reproduction does occur in most

unicellular eukaryotes. Diploid cells undergo
reduction division (meiosis) to make haploid
gametes or simply haploid gamete nuclei.
The haploid gametes or nuclei fuse to make a
diploid stage or a zygote, but there are no
embryos.
The life cycles vary greatly in how much time is
spent in haploid and diploid stages, and in the
presence or absence of asexual multiplication.

Paramecia reproduce only by binary fission

Although all unicellular eukaryotes reproduce
asexually, and some are apparently exclusively
asexual, the widespread occurrence of sex
testifies to its importance as a means of genetic
recombination.
Gamete nuclei, or pronuclei, which fuse in
fertilization to restore the diploid number of
chromosomes, are usually borne in special
gametic cells.
When gametes all look alike, they are called
isogametes, but most species have two
dissimilar types, or anisogametes.
across kineties (ciliary rows) but have certain
forms of sexual phenomena called conjugation
and autogamy.
In binary fission the micronucleus divides
mitotically into two daughter micronuclei,
which move to opposite ends of the cell. The
macronucleus elongates and divides
amitotically.
Conjugation occurs at intervals in ciliates.
Conjugation is the temporary union of two
individuals to exchange chromosomal material
(Figure 11.26).
During the union the macronucleus
disintegrates and the micronucleus of each
individual undergoes meiosis, giving rise to four
haploid micronuclei, three of which degenerate
(Figure 11.26A–C).
The remaining micronucleus then divides into
two haploid pronuclei, one of which is
exchanged with the other conjugant.
The pronuclei fuse to restore the diploid
number of chromosomes, followed by several
more nuclear events detailed in Figure 11.26.
Following this complicated process, the
organisms may continue to reproduce by binary
fission without conjugation.
The result of conjugation is similar to that of
zygote formation, for each exconjugant contains
hereditary material from two individuals.
Sexual reproduction permits gene
recombinations, thus increasing genetic
variation in the population.

The cell multiplication process that produces

more individuals in unicellular eukaryotes is
called fission.
he most common type of fission is binary, in
which two essentially identical individuals
result.
When a progeny cell is considerably smaller than
the parent and then grows to adult size, the
process is called budding.
Multiple fission, or schizogony, is common
among the Apicomplexa and some amebas.

LIFE CYCLE OF PLASMODIUM

Life cycle of Plasmodium vivax, one of the

protozoa (class Coccidia) that causes malaria in
humans.
 A. Sexual cycle produces sporozoites in a more likely that the absence of mitochondria is a
mosquito. Meiosis occurs just after zygote secondary loss, not a primary absence.
formation (zygotic meiosis).
B. Sporozoites infect a person and reproduce
asexually, first in liver cells and then in red blood
cells. Malaria is spread by Anopheles mosquito,
which ingests gametocytes (sex cells) along with
human blood, then, when biting another victim,
leaves sporozoites in new wound.

Part 3: Major Eukaryotic Clades

Giardia, a diplomonad, is a well-studied parasite.

Some species inhabit the human digestive tract,
but others occur in birds or amphibians. It is
often asymptomatic but may cause a rather
discomfiting, but not fatal, diarrhea. Cysts are
passed in the feces, and new hosts are infected
by ingestion of cysts, often in contaminated
water.

PARABASALIDS

Members of this phylum have a stiffening rod

composed of microtubules, the axostyle, that
extends along the longitudinal axis of their body.
Cladogram showing the major eukaryotic clades; Parabasalids, traditionally part of the class
in many cases the order of branching remains to Parabasalea, possess a modified region of the
be determined. Opisthokonta is a very large Golgi complex called a parabasal body,
clade comprising choanoflagellates, fungi, and hydrogenosomes, a characteristic flagellated
multicellular animals. The terms “SAR” and stage, and a unique type of mitosis.
“RAS” are interchangeable; SAR represents the
first letter of the names Stramenopiles,
Alveolates, and Rhizaria, whereas RAS
represents the same taxa listed in the reverse
order.

PHYLUM RETORTAMONADA AND THE DIPLOMONADS

This phylum is divided into two exclusive clades:

retortamonads and diplomonads.
Retortamonads include commensal and
parasitic unicells, such as Chilomastix and Much of the work on parabasalid structure has
Retortamonas. They lack mitochondria and been done on species of Trichomonas, a disease-
Golgi bodies. causing organism for humans and other animals.
Diplomonads, once a subgroup of Some trichomonads are of medical or veterinary
retortamonads, also lack mitochondria, and importance.
were likewise proposed to lie outside the clade
PHYLUM EUGLENOZOA
of mitochondria-containing eukaryotes.
However, work showing that mitochondrial The Euglenozoa is generally considered a
genes occur in the cell nucleus1 makes it much monophyletic group, based on the shared
persistence of the nucleoli during mitosis, and
the presence of discoid mitochondrial cristae.
Members of this phylum have a series of
longitudinal microtubules just beneath the
plasma membrane that help to stiffen the
membrane into a pellicle.
The phylum is divided into two subphyla, the
Euglenida and the Kinetoplasta.
Euglenids have chloroplasts with chlorophyll b.
These chloroplasts are surrounded by a double
membrane and are likely to have arisen via
secondary endosymbiosis.
Euglena viridis is a representative flagellate
commonly studied in introductory zoology
courses.
Its natural habitat is freshwater streams and
ponds where there is considerable vegetation.
The organisms are spindle shaped and about
60µm long, but some species of Euglena are
smaller and some larger (E. oxyuris is 500µm
long).
Just beneath the outer membrane of Euglena are
proteinaceous strips and microtubules that form
a pellicle.
In Euglena the pellicle is flexible enough to
permit bending, but in other euglenids it may be
more rigid.
Nutrition of Euglena is normally autotrophic
(holophytic), but if kept in the dark the organism
uses saprozoic nutrition, absorbing nutrients
through its plasma membrane.
Mutants of Euglena can be produced that have
permanently lost their photosynthetic ability.
Although Euglena does not ingest solid food,
some euglenids are phagotrophic.
Peranema has a cytostome that opens alongside
its flagellar reservoir.
Encyst – become enclosed in a cyst
Kinetoplastans are named for the presence of a
unique organelle, the kinetoplast. This modified
mitochondrion, associated with a kinetosome,
carries large discs of DNA.
Kinetoplastans are all parasites of plants or
animals.
Some are nonpathogenic, but others produce
severe diseases in humans and domestic
animals.
Trypanosoma brucei gambiense and T. brucei
rhodesiense cause African sleeping sickness in
humans
T. brucei brucei causes a related disease in
domestic animals.
Trypanosomes are transmitted by tsetse flies
(Glossina spp.)
Trypanosoma b. rhodesiense, the more virulent
of the sleeping sickness trypanosomes, and T. b.
brucei have natural reservoirs (antelope and
other wild mammals) that are apparently not
harmed by the parasites.
Trypanosoma cruzi causes Chagas’ disease in
humans in Central America and South America.
It is transmitted by “kissing bugs” (Triatominae),
a name arising from the bug’s habit of biting its
sleeping victim on the face.
Acute Chagas’ disease is most common and
severe among children less than five years old,
while the chronic disease occurs most often in
adults.
Symptoms are primarily a result of central and
peripheral nervous dysfunction.
 Symbiotic Ciliates
CLADE ALVEOLATA
In the Ciliophora, the alveoli produce pellicles
Dinoflagellata, a group of armored flagellates,
the alveoli produce thecal plates
in the Apicomplexa, containing intracellular
parasitic species previously called sporozoans,
the alveoli have structural functions.
PHYLIUM CILIOPHORA
Ciliates are so named because the body surface
is covered with cilia that beat in a coordinated,
rhythmical manner.
The arrangement of cilia varies within the
phylum and some ciliates lack cilia as adults,
although cilia occur at other stages in the life
cycle.
In general, ciliates are larger than most other
unicellular eukaryotes, but they range from
10µm to 3 mm in length.
Ciliates are very structurally complex, exhibiting
a wide range of specializations.
In other forms there may be fused cilia forming
stiffened tufts called cirri, often used in
locomotion by the creeping ciliates.
Many symbiotic ciliates live as commensals, but
some can be harmful to their hosts.
Balantidium coli lives in the large intestine of
humans, pigs, rats, and many other mammals.
Some strains seem to be host-specific, so the
organism is not easily transmitted from one
species to another.
Transmission is by fecal contamination of food
or water.
Usually the organisms are not pathogenic, but in
humans they sometimes invade the intestinal
lining and cause a dysentery similar to that
caused by Entamoeba histolytica.
The disease can be serious and even fatal.
Infections are common in parts of Europe, Asia,
and Africa but are rare in the United States.
Entodinium belongs to a group that has very
complex structure and lives in the digestive tract
of ruminants (cow, sheep, deer, giraffe, etc.),
sometimes in great abundance.
Nyctotherus inhabits the colon of frogs and
toads.
 In aquarium and wild freshwater fishes,
Ichthyophthirius causes a disease known to
many fish culturists as “ick.” Untreated, it can
cause much loss of exotic fishes.
Free-living Ciliates
Stentor (Gr. herald with a loud voice) – trumpet
shaped and solitary, with a beadshaped
macronucleus
Vorticella (L. dim. of vortex, a whirlpool) – bell
shaped and attached by a contractile stalk
Euplotes (Gr. eu, true, good, + ploter, swimmer)
– with a flattened body and groups of fused cilia
(cirri) that function as legs
Paramecia are usually abundant in ponds or
sluggish streams containing aquatic plants and
decaying organic matter. Slipper-shaped.
PHYLUM DINOFLAGELLATA
Ecologically, some species are among the most
important primary producers in marine
environments.
They commonly have two flagella, one
equatorial and one longitudinal, each borne at
least partially in grooves on the body.
The body may be naked or covered by cellulose
plates or valves.
Many species can ingest prey through a mouth
region between the plates near the posterior
area of the body.

Ceratium bears plates and is

both autotrophic and phagotrophic
Ceratium has a thick covering
with long spines, into which the body
PHYLUM APICOMPLEXA
extends, but it can catch food with
posterior pseudopodia and ingest it
between the flexible plates in the
posterior groove.
Gymnodinium bears no cellulose
plates.
Almost all apicomplexans are endoparasites,
and their hosts include many animal phyla.
The presence of a certain combination of
organelles, the apical complex, distinguishes
this phylum.
Locomotor organelles are less obvious in this
group than in other protozoa.
Pseudopodia occur in some intracellular stages,
and gametes of some species are flagellated.
The life cycle usually includes both asexual and
sexual reproduction, and sometimes an
invertebrate intermediate host.
At some point in the life cycle, the organisms
develop a spore (oocyst), which is infective for
the next host and is often protected by a
resistant coat.

Noctiluca, a colorless
dinoflagellate, is a voracious
predator and has a long, motile
tentacle, near the base of which
its single, short flagellum
emerges. Toxoplasma – causes toxoplasmosis, a disease
Noctiluca is one of many affecting cats and humans
marine organisms that can Plasmodium – the organism that causes
produce light (bioluminescence). malaria.
Eimeria – generally affects birds
Plasmodium: The Malarial Organism

The best-known coccidians are Plasmodium

spp., causative organisms of the most
important infectious disease of humans:
malaria.
Malaria is a very serious disease, difficult to
control and widespread, particularly in tropical
and subtropical countries.
Four species of Plasmodium infect humans: P.
falciparum, P. vivax, P. malariae, and P. ovale.
Although each species produces its own
peculiar clinical picture, all four have similar
cycles of development in their hosts.

Chlamydomonas is a unicellular biflagellate cell.

Colony formation in green algae occurs when
the products of cell division maintain
cytoplasmic bridges between cells and cells sit
within an extracellular matrix (ECM).
True multicellularity requires a division of labor
among cells: some cells become the germ line
and others become the somatic line.
The genetic changes necessary to set up this
division have been elucidated in the green algal
CLADE PLANTAE lineage that produced a multicellular species,
Volvox carteri.
The clade Plantae comprises three
A colonial body plan is visible in Gonium,
photosynthetic lineages: glaucophytes,
Eudorina, and Pandorina (Figure 11.33), but
rhodophytes (red algae), and Viridiplantae.
Volvox was one of nine green algal lineages in
Plantae is sometimes called Archaeplastida in
which multicellularity evolved independently.
reference to the ancient primary symbiosis with
a cyanobacterium that produced the
chloroplasts of photosynthetic eukaryotes.
Viridiplantae contains the unicellular, colonial,
and multicellular green algae, which were once
in phylum Chlorophyta, as well as the
nonvascular and vascular plants.
The phylum Chlorophyta is a non-monophyletic
group containing only the green algae.
We now know that the vascular and nonvascular
plants (jointly called “land” plants) shared a
common ancestor with the green algae.
AMOEBOZOA
Amoebozoans include naked and testate
amebas, as well as amebas with flagellated
stages in the life cycle.
Amoebozoans typically have branched tubular
mitochondrial cristae, but this feature is not
unique to the group.
Ameboid forms within the group include the
fascinating plasmodial and cellular slime molds
in the subgroup Mycetozoa (e.g., Physarum and
Dictyostelium, respectively).
Amoebozoa also houses entozoic amebas—
those that live inside humans or other animals.
Entamoeba histolytica is an important human
parasite. It lives in the large intestine and on
occasion can invade the intestinal wall by
secreting enzymes that attack the intestinal
lining.
If this occurs, a serious and sometimes fatal
amebic dysentery may result. The organisms
may be carried by the blood to the liver and
other organs and cause abscesses there.
Many infected persons show few or no
symptoms but are carriers, passing cysts in their
feces.
Diagnosis is complicated by the existence of a
nonpathogenic species, E. dispar, which is
morphologically identical to E. histolytica.
Infection is spread by contaminated water or
food containing cysts.
E. histolytica occurs throughout the world, but
clinical amebiasis is most prevalent in tropical
and subtropical areas.
Other species of Entamoeba found in humans
are E. coli in the intestine and E. gingivalis in the
mouth. Neither of these species is known to
cause disease.