Euglena
Dennis E Buetow, University of Illinois, Urbana, Illinois, USA
Euglena is a genus of single-celled, free-living micro-
organisms that show both plant- and animal-like
characteristics. Members of the genus are found widely
in nature and mainly in fresh waters. Most are aerobic
and use photosynthesis or many organic compounds as
interchangeable sources of carbon and energy. The most
studied species is the easily grown E. gracilis which, in spite
of its relatively rigid surface, can be subfractionated to
yield nuclei, mitochondria, chloroplasts, flagella and pel-
licles. Subfractions show some unusual characteristics:
The nuclear envelope remains intact throughout mitosis
and chromosomes are permanently condensed at all
phases of the cell cycle, the mitochondrial respiratory
chain has two terminal oxidases, and chloroplasts have
three membranes reflecting their endosymbiotic evo-
lutionary history. Various Euglena and some of their
constituent molecules are useful for environmental bio-
mediation and potentially useful biomedically.
Introduction
Antoni van Leeuwenhoek in 1674 was the rst to describe
a microorganism which was green in the middle. This
organism later was named Euglena viridis by CG Ehrenberg
(1830) who started the study of the genus Euglena. More
than 250 Euglena species have been described. The actual
number is not known, however, because the size and shape
of a cell in a given species can vary greatly depending on
nutritional status, phase of the growth cycle (lag, exponen-
tial or stationary) and environmental factors such as pH and
the quantity and quality of light available. Therefore, some
species described may only be size variants of other better-
dened species. In any case, the single-celled organisms
comprising the genus Euglena are structurally complex
and show both plant- and animal-like characteristics. All
ELS subject area: Microbiology
How to cite:
Buetow, Dennis E (April 2011) Euglena. In: Encyclopedia of Life
Sciences (ELS). John Wiley & Sons, Ltd: Chichester.
DOI: 10.1002/9780470015902.a0001964.pub3
ENCYCLOPEDIA OF LIFE SCIENCES & 2011, John Wiley & Sons, Ltd. www.els.net
Advanced article
Article Contents
. Introduction
. Description and Taxonomy
. Reasons for Popularity
. Significance for Research
. Impact on Human Welfare
Online posting date: 15th April 2011
known Euglena species are asexual. A very large literature
exists on the morphology, physiology and biochemistry of
Euglena, but much of the information comes from only a
few species, with the easily grown Euglena gracilis being the
most studied (Buetow, 19681989). See also: Ehrenberg,
Christian Gottfried; Leeuwenhoek, Antoni van
Description and Taxonomy
Morphology
Euglena cells range in size from 12 mm long  5 mm wide
for E. minuta to 530 mm  40 mm for E. oxyuris and vary
in shape from almost spherical to nearly cylindrical.
The largest numbers are spindle shaped. Locomotion is
accomplished by the beating of the long agellum.
Metaboly, or euglenoid movement, also occurs in most if
not all species and, therefore, the overall shape of the cell
may change at any time. Figure 1 (Leedale, 1982) depicts
that a Euglena cell of the E. gracilis type showing its
main organelles. E. gracilis cells are approximately 50 mm
long  10 mm wide and spindle-shaped. The majority of
Euglena species are green due to the presence of chloro-
plasts containing chlorophylls a and b. The chloroplasts of
E. gracillis are labile and easily lost being readily bleached
by a variety of agents including antibiotics and tempera-
tures of 348C or above. About a dozen species are coloured
by red granules. When the granules are concentrated at its
centre, the cell appears green because of the peripheral
arrangements of its chloroplasts. When the granules are
dispersed, the cell appears red. The surface of most, per-
haps all, Euglena cells is coated with a mucilage that is
secreted from muciferous bodies via canals to the outside.
May species encyst especially under unfavourable envir-
onmental conditions. See also: Cilia and Flagella
Ecology
Members of the genus Euglena are found widely in nature.
The species are free living and inhabit freshwater pools,
ponds and lakes. Apparently, there are no saltwater
forms, though some Euglena occasionally are found in
marine sediments and some survive in up to 40% seawater.
Euglena species generally are aerobic, but some tolerate
anaerobic conditions. Various species are indicators of
1
 Euglena
Figure 1 A Euglena cell of the Euglena gracilis type showing the main
organelles. From Leedale (1982).
organically polluted water. E. gracilis survives high levels
of highly energetic ionising radiation (Hayashi et al., 2004).
An unusual degree of adaptability to diverse environments
characterises the genus, with species found in vegetable
cannery and citrus waste-lagoons, in raw sewage, in water
in tree holes, on snow and in high mountain lakes, on the
bark of the honey locust tree, in alkaline marshes and in
acid coal-mine water with a pH as low as 0.9 (Lacky, 1968).
E. gracilis survives temperatures from about 18C to 388C
and pHs ranging from 2.3 to 11. The most thermotolerant
(up to 408C) chloroplast-bearing Euglena strain known
lives in acidic hot mud pools in a volcanic area in Costa
Rica (Sittenfeld et al., 2002). See also: Protozoan Ecology
Nutrition
As a group, Euglena are able to use photosynthesis and
heterotrophic oxidative assimilation as interchangeable
and apparently equivalent sources of carbon and energy.
The degree of interchangeability, however, can vary from
2 ENCYCLOPEDIA OF LIFE SCIENCES & 2011, John Wiley & Sons, Ltd. www.els.net
species to species and even from strain to strain. At one
extreme is E. pisciformis, an obligate phototroph, and at
the other are the articially bleached strains of Euglena,
which are obligate heterotrophs. The Vischer strain grows
more rapidly in light than on any of several organic
substrates. E. gracilis strain Z and variety bacillaris can
grow on a variety of media. Minimally, they require several
inorganic salts as sources of phosphorus, sulfur, nitrogen
and minerals, vitamins B1 and B12 and white light or any
one of the many organic compounds as a source of carbon
(Cook, 1968).
Taxonomy
How to classify Euglena has been a long-standing problem
(Linton et al., 2010). The genus has been claimed as
photosynthetic protozoa by zoologists and as algae by
botanists. However, inclusion of the genus among the
algae or the protozoa may no longer be tenable. Molecular
phylogeny based on the nucleotide sequences of genes
encoding small-subunit ribosomal ribonucleic acids (RNAs)
(Sogin et al., 1986) shows E. gracilis diverging from the
main eukaryotic line far before a period of massive
evolutionary radiation that gave rise to the algae, plants,
animals and fungi. The free-living organisms of the genus
Euglena have been placed in the Phylum Euglenozoa, which
interestingly also includes the extant parasitic trypanosomes
(Von der Heyden et al., 2004; Ahmadinejad et al., 2007)
See also: Protozoan Taxonomy and Systematics
Reasons for Popularity
E. gracilis was among the rst of the photosynthetic
eukaryotic microorganisms domesticated for laboratory
use. It is easy to grow, with various strains showing gen-
eration times of 10 to about 25 h at 258C. The biochemical
and physiological exibility of this species, together with a
morphological exibility that is readily visualised via light
and electron microscopy, contribute to its popularity. It is
amenable to single-cell studies as well as studies using large-
scale fermenter-size cultures. The cell cycle of E. gracilis can
be studied in mass cultures synchronised for cell division
through the use of alternating light and dark cycles or heat
and cold shock. This species is a prominent example that
circadian and infradian rhythms exist and can be investi-
gated readily in single-celled organisms. Further, in spite of
its relatively rigid pellicle, E. gracilis can be subfractionated
to yield pure preparations of nuclei, chloroplasts, mito-
chondria, agella and pellicles. See also: Cell Cycle;
Tetrahymena
Significance for Research
Studies on the genus Euglena, and especially E. gracilis,
have been signicant for understanding the biochemistry
and molecular biology of nuclei and subcellular organelles.

Also, some unusual features of these subcellular particles
in Euglena distinguish them from their counterparts in
higher eukaryotes and, so, have furthered our knowledge
of evolutionary diversity.
Nucleus
The Euglena nucleus has been isolated and its ultrastucture
extensively investigated. The number of chromosomes
varies with the species, e.g., E. gracilis has 45 while the
higher ploidy E. spirogyra has 86. The chromosomes,
unlike those in higher cells, appear permanently condensed
at all phases of the cell cycle (Leedale, 1967). Even so,
chromatin can be isolated from interphase Euglena nuclei
with over 60% being highly condensed heterochromatin
and the rest being the less condensed, transcriptionally
active euchromatin. The usual histones are present.
Nuclear division in Euglena is by mitosis, but a peculiar one
in that the nuclear envelope persists throughout the entire
process; the nucleolus (also called an endosome, Figure 1)
also remains intact, elongates along the division axis and
divides into two daughter nucleoli; and the chromosomes
are arranged parallel to the division axis at metaphase.
Some species appear to have several nucleoli, and these
fuse into a single body at mitosis. Many Euglena nuclear
mRNAs, are subject to trans-splicing rather than the more
common cis-splicing found in higher eukaryotes (Tessier
et al., 1991; Frantz et al., 2000). See also: Eukaryotic
Chromosomes; Nucleolus
Mitochondrion
The mitochondrial system of E. gracilis and apparently
of other Euglena species is usually a single reticulum that
ramies throughout the cell (Figure 1). The degree of
branching and the thickness of the branches, however, may
vary under dierent nutritional conditions and during
dierent phases of the cell cycle. The outer mitochondrial
membrane is strongly undulated and the cristae, unlike
those in most eukaryotes but like those in trypanosomes,
are discoid (previously called at) and constricted at their
bases. Also, the mitochondrial succinate dehydrogenase
enzyme is encoded by a unique split and rearranged nuclear
gene in E. gracilis as it is in trypanosomes (Gawryluk and
Gray, 2009).
The mitochondrion of E. gracilis shares with all mito-
chondria a common basic pattern of oxidative phos-
phorylation and participation in intermediary metabolism
(Buetow, 1989; Kitaoka et al., 1989). It also shows some
apparently unique characteristics. For example, its tri-
carboxylic acid (TCA) cycle in overall function is similar to
that in other mitochondria but contains novel NADP+-
dependent 2-oxoglutarate decarboxylase (OGD) and
pyruvate dehydrogenase (PD) enzymes. Both function as
single enzymes rather than as part of a complex of enzymes
and are dependent on NADP+ rather than NAD. The
Euglena PD is oxygen-sensitive and functions as a regu-
latory enzyme depending on the concentration of oxygen.
ENCYCLOPEDIA OF LIFE SCIENCES & 2011, John Wiley & Sons, Ltd. www.els.net
Euglena
At a normal level of oxygen (21%), it functions in the
TCA cycle, while at limiting oxygen (essentially anaerobic)
conditions, it functions in wax ester fermentation. No
adenosine triphosphate (ATP) is lost by this unique process
of wax fermentation. Therefore, the ATP generated by
glycolysis remains as a net gain and apparently allows the
cells to survive anaerobiosis. The Euglena OGD converts
2-oxoglutarate to succinate semialdehyde (SSA) in con-
trast to the 1-oxoglutarate dehydrogenase which provides
for the formation of succinyl-coenzyme A in the TCA cycle
of other mitochondria. The SSA of Euglena also is part of a
unique g-aminobutyric acid (GABA) shunt similar to the
GABA shunt found in mammalian brain mitochondria. In
addition to oxidative phosphorylation, ATP is also gen-
erated in the Euglena mitochondrion by adenylate kinase
and by a catalatic-type activity (but not catalase-driven)
that converts hydrogen peroxide to water and oxygen with
the energy released used to drive the formation of ATP. The
respiratory chain contains two terminal oxidases and sep-
arate entry points for oxidation of NADH, succinate and
lactate. The alternative oxidase is inducible, highly resist-
ant to cyanide and contains haem B as do bacterial quinol
oxidases (Devars et al., 1998). Glyoxylate cycle enzymes
are present in mitochondria rather than in glyoxysomes
(Ono et al., 2003). The Euglena mitochondrion unites
biochemical properties of aerobic and anaerobic mito-
chondria and of hydrogenosomes (Homeister et al.,
2004). See also: Citric Acid Cycle; Mitochondria: Structure
and Role in Respiration; Oxidative Phosphorylation
Euglena mitochondrial deoxyribonucleic acid (DNA)
has a molecular mass of 40  106 Da and a low (25%) G+C
content and appears to exist as a complex mixture of
circular and heterogeneous linear molecules similar to the
situation encountered with higher plant mitochondrial
genomes (Buetow, 1989).
Chloroplast
Chloroplasts in Euglena species range from small pyrenoid-
less discs to large plates or complexes with pyrenoids,
photosynthetic lamellae typically with three thylakoids
and a matrix (or stroma) containing 70S ribosomes. In
contrast to green algal and higher plant chloroplasts, which
are surrounded by two membranes, Euglena chloroplasts
are surrounded by three membranes. Thus, the latter
organelles evolved from a secondary endosymbiosis with a
eukaryotic symbiont rather than a primary endosymbiosis
with a prokaryotic symbiont (Gibbs, 1978; Archibald and
Keeling, 2002). In green algae and higher plant chloro-
plasts, protein import is posttranslational. In Euglena,
the presence of an extra chloroplast membrane leads to
an unusual import pathway with many nuclear-encoded
chloroplast proteins being transported rst to the endo-
plasmic reticulum then to the Golgi and then to the chloro-
plast (Sulli and Schwartzbach, 1996; Slavikova et al., 2005).
Light is indispensable for plastid development in
E. gracilis (Schi and Schwartzbach, 1982). During hetero-
trophic growth in darkness, plastid development is stopped
3
 Euglena
at the stage of a small proplastid. On exposure to light, the
proplastid is induced to develop into a chloroplast. Thus,
chloroplast development is easily studied in E. gracilis by
rst growing the cells in the dark and then simply exposing
them to white light. Therefore, more research has been
devoted to the chloroplasts of E. gracilis than to any other
component of a Euglena cell. These chloroplasts are rela-
tively streamlined to function mainly as photosynthetic
units. Unlike those in higher plants, Euglena chloroplasts,
for example, do not store starch, do not activate or reduce
sulfate, and do not contain an NADP+-dependent malate
dehydrogenase. Sulfur metabolism and the dehydrogenase
are located in mitochondria in Euglena (Saidha et al., 1988;
Patton et al., 2008). See also: Photosynthesis
Chloroplast DNA in E. gracilis is A+T-rich (about
75%) and has a molecular mass of 92  106 Da. There
are about 2002800 DNA molecules in the 610 chloro-
plasts in a Euglena cell with the exact number depending
on the condition of growth and the phase of the growth
cycle. The chloroplast DNA has been completely seque-
nced (Hallick et al., 1993), this circular DNA is 143 170 bp
in size, counting only one copy of a 54-bp tandem
repeat that is present in variable copy number. In contrast
to the 120130 genes in land-plant chloroplast genomes,
there are only 103 in Euglena. Introns account for
38.3% of the total DNA content and are the most
numerous of any organelle genome known. Included are
twintrons (introns-within-introns).
Impact on Human Welfare
E. gracilis requires low levels of vitamin B12 for growth.
Therefore, the rst use of this organism for medical pur-
poses was in assays to determine B12 levels in biological
uids. E. gracilis shows promise as a single-cell source of
protein (Hosoya and Kitaoka, 1977; Chae et al., 2006) and
as an antihypertensive agent (Murakami, 1985), provides
an ecient and inexpensive system for purication and
reclamation of farm and urban wastes (Waygood et al.,
1980), can serve as a bioreactor for the production of wax
esters (Tani et al., 1987), is highly resistant to heavy metals
and can be used to indicate the health of the environment
(Rodriguez-Zavala et al., 2007), is suitable for remediating
water contaminated by radioactive technium (Ishii and
Uchida, 2006) and is promising for commercial production
of g-tocopherol which is used as a dietary supplement for
humans, as a food preservative, and in the manufacture
of cosmetics and for the fortication of animal feed (Tani
and Tsumura, 1989; Ogbonna, 2009). Paramyton, a major
glucan in E. gracilis, protects mammalian liver from injury
by carbon tetrachloride and inhibits dermatitis-like skin
lesions (Sugiyama et al., 2009, 2010).
Plants, transformed with an unusual desaturase gene
from Euglena, produce essential fatty acids normally
obtained only from sh (Green, 2004). E. ignobilis, based
on mobility and morphological changes, can be used to
access water quality (Gupta and Agrawal, 2005). E. viridis
4 ENCYCLOPEDIA OF LIFE SCIENCES & 2011, John Wiley & Sons, Ltd. www.els.net
is cost eective for degrading swine manure in polluted
agro-wastewater (de Godos et al., 2010). E. sanguinea
contains a toxin, euglenophycin, which has herbicidal
and anticancer activities (Zimba et al., 2010). See also:
Bioremediation
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Further Reading
Ciugulea I and Triemer RE (2010) A Color Atlas of Photosynthetic
Euglenoids. East Lansing: Michigan State University Press.
Gojdics M (1953) The Genus Euglena. Madison: The University of
Wisconson Press.
Kitaoka S (1989) Euglena-Physiology and Biochemistry. Tokyo:
Gakkai Shuppan Center. [In Japanese].
Leedale GF (1971) The Euglenoids. Oxford Biology Readers.
London: Oxford University Press.
Leedale GF (2000) Euglenozoa. In: Lee JJ, Leedale GF and
Bradbury P (eds) An Illustrated Guide to the Protozoa, 2nd edn,
vol. II, pp. 11351185. Lawrence, KS: Allen Press, Inc.
Sleigh M (1989) Protozoa and Other Protists. London: Edward
Arnold.
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