Slime Moulds

Secondary article
Article Contents

Richard L Blanton, Texas Tech University, Lubbock, Texas, USA

. Brief Description and Characterization

Slime moulds are organisms that feed and grow like protozoa but reproduce like fungi.
Their unusual name belies their importance as experimental research organisms and their
potential to provide insights into the evolution of multicellularity.

Brief Description and Characterization

Slime moulds are eukaryotic microorganisms with life
cycles that alternate between a protozoan-like phagotrophic stage, responsible for feeding and growth, and a
mould-like stage, responsible for reproduction. The
mould-like stage involves the formation of a fruiting body,
which encloses or bears spores. The slime mould lifestyle or
habit is also found in bacteria (most notably the
myxobacteria) and a ciliated protozoan (Sorogena stoianovitchae, a true member of the phylum Ciliophora). In
some slime mould groups the transition from the trophic to
the reproductive stage involves the aggregation of individual cells to form a multicellular structure that then
produces the fruiting body. This, too, is accomplished by
myxobacteria and Sorogena. Most of the organisms that
have been called slime moulds have phagotrophic stages
that are either amoebae or coenocytic structures called
plasmodia. Closer examination of the amoebal stages,
patterns of aggregation and mechanisms of fruiting body
development indicate that the slime mould habit arose in
several unrelated amoebal groups. Therefore, the slime
mould habit, with or without aggregation, is a fascinating
example of convergent evolution but is not useful as a
criterion for the phylogenetic classication of organisms.
The cellular slime mould Dicytostelium discoideum
provides an excellent example of the life cycle of a slime
mould. The feeding and multiplication stage consists of
unicellular amoebae, which feed on bacteria in the moist
humus layer of forest soil. Upon the exhaustion of the food
source, the amoebae aggregate to form a multicellular
structure called a pseudoplasmodium or slug (because of
its supercial resemblance to a garden slug). The cells in the
slug no longer feed or divide. Instead, the slug is a
migratory stage that serves to move the population of
amoebae to a more favourable environment for the
dispersal of spores. In the forest, this would be atop the
leaf litter. Once it has reached appropriate conditions, the
slug forms an aerial fruiting body consisting of a tapering
stalk with a mass of spores at its apex. These spores are
dispersed by insects and other means. A spore will
germinate to yield one amoeba, thereby beginning the
cycle once again.
Four groups of microorganisms have been called slime
moulds: the plasmodial slime moulds, the cellular slime
moulds, the endoparasitic slime moulds (the Plasmodio-

. Place in Overall Taxonomic Scheme

. Major Subtaxa and Well-known Species
. Phylogenetic and Evolutionary Considerations

phoromycetes or Plasmodiophorales (Karling, 1968;

Braselton, 1995)), and the net slime moulds (the Labyrinthulomycota or Labyrinthulales (Porter, 1990)). The
latter two groups do not have the true slime mould habit:
their trophic stage is absorptive, not ingestive. This article
is concerned with the plasmodial and cellular slime moulds.
In addition, it includes a more recently discovered group of
fruiting protists, the protostelids, which appear to be
related to the plasmodial and cellular slime moulds.
Slime moulds overall are cosmopolitan in distribution,
but individual species and groups may have highly
specialized habitats and/or limited distribution. They are
secondary decomposers, feeding on the cells of bacteria
and yeast and the conidia and spores of fungi (and, in some
cases, the amoebae of other slime moulds). The larger
species of plasmodial slime moulds can be readily seen with
the unaided eye on fallen leaves, rotting logs and forest
litter; some species commonly appear on lawns or garden
mulch. Other slime moulds can be isolated in the
laboratory from similar substrates, as well as from tree
bark, animal dung, decaying aerial plant parts, humus and
soil.
Slime moulds have no known economic importance.
They are not known to be pathogenic to humans, animals
or plants. Some plasmodial slime moulds can temporarily
discolour lawns with their fruiting bodies. D. discoideum, a
cellular slime mould, and Physarum polycephalum, a
plasmodial slime mould, have great value as experimental
research organisms in biomedical research. D. discoideum
may have economic value in the future as a microbial
expression system for introduced genes.

Place in Overall Taxonomic Scheme

Mycologists discovered slime moulds and have described
most of the species and higher taxa. Hence, slime moulds
have traditionally been classied as fungi and their group
name endings reect this classication (e.g. Myxomycetes
for the plasmodial slime moulds; Acrasiomycetes for the
cellular slime moulds). Their distinctiveness from the
lamentous fungi was usually recognized by placing the
slime moulds in a separate high-level grouping. For
instance, in a kingdom Fungi, the subkingdoms would

ENCYCLOPEDIA OF LIFE SCIENCES 2001, John Wiley & Sons, Ltd. www.els.net

Slime Moulds

include Eumycota (true fungi) and Myxomycota (slime

moulds).
From the time of their discovery in the late nineteenth
century, the animal or protozoan aliation of the slime
moulds was recognized by the mycologist Anton deBary,
as evidenced by the sux of the name he proposed for
them, the Mycetozoa. Today, most mycologists and
protozoologists would classify the slime moulds in the
kingdom Protista (or Protozoa). However, the slime
moulds would not occupy a single taxon because they
include a number of unrelated organisms.
There is strong evidence that the plasmodial slime
moulds, most of the protostelids and the subgroup of
cellular slime moulds called the dictyostelids are related
and can be grouped together as the Eumycetozoa. The
acrasid cellular slime moulds includes four groups of
unrelated organisms that are classied together solely on
the basis of being aggregative amoebae displaying the slime
mould habit. Each of these four groups would more
properly be classied together with their closest nonfruiting relatives. For instance, the fruiting genera Acrasis and
Pocheina have been grouped together with nonfruiting
genera such as Naegleria and Tetramitus in the Heterolobosea, the rst taxon to include both fruiting and
nonfruiting amoeboid organisms (Page and Blanton,
1985). Other members of the acrasid cellular slime moulds
and some of the noneumycetozoan protostelids have yet to
nd a taxonomic home.

Major Subtaxa and Well-known Species

The Eumycetozoa
The Eumycetozoa includes three groups: the plasmodial
slime moulds (also called the true slime moulds, the
acellular slime moulds, the myxomycetes or the myxogastria); the dictyostelid cellular slime moulds; and the
protostelids. Upon initial examination, the plasmodial
and dictyostelid cellular slime moulds would appear to be
unrelated. However, the discovery of the protostelids
provided a common group from which both of these
groups apparently evolved. As will be discussed below,
there is increasing molecular systematic evidence for the
Eumycetozoa.
Protostelids
The protostelids are structurally the simplest slime moulds.
The nearly 40 described species are microscopic, with the
exception of Ceratiomyxa. The trophic stage is an amoeba,
amoeboagellate (with two agella) or small plasmodium.
Single cells or plasmodia produce a stalked fruiting body
consisting of an acellular stalk and usually a single spore.
Some species produce fruiting bodies with more than one
spore as a result of mitosis occurring during fruiting body
2

formation. Amoebae can form cysts in most species. Sexual

processes are suggested in some species, but have not been
proven genetically. Ultrastructural studies of mitosis and
the agellar apparatus provide strong evidence for the
relatedness of the agellated protostelids and the plasmodial slime moulds (Spiegel et al., 1995). The similarities
between the two groups can be further illustrated by the
reclassication of the plasmodial slime mould Ceratiomyxa as a protostelid and the transfer of the protostelid
Cavostelium bisporum to the plasmodial slime mould genus
Echinostelium. The protostelids have been relatively little
studied and hence cannot really be said to have well-known
species; Planoprotostelium aurantium has been used in
recent sequence analysis studies.
Plasmodial slime moulds
The plasmodial slime moulds number over 50 genera and
500 species organized into three subclasses, six orders and
13 families. The fruiting body of a plasmodial slime mould
is called a sporocarp (the spores are contained within a
common wall). Sporocarps show a diversity of forms,
colours and sizes (ranging from 0.05 mm to over 5 cm).
They may be sessile or stalked; the stalks are acellular and
do not have a stalk tube as in the dictyostelid cellular slime
moulds. The sporangial wall encloses a mass of spores,
which in many species have ornamented walls. The spores
may be interspersed within a network of noncellular
threads called capillitia, which may also be ornamented
or sculpted. The spores germinate to yield 14 amoebae or
amoeboagellates (called swarm cells); these cells interconvert, depending upon moisture levels and possibly
other factors. The swarm cell has two anterior whiplash
agella, one long and directed anteriorly and the other very
short and recurved. The cells swim with a rotating
movement and alternate periods of swimming with periods
of crawling by amoeboid movements. The ultrastructure of
the agellar apparatus of the swarm cell is conserved across
the group, and its similarity to that of the agellated
protostelids provides strong evidence for the relatedness of
the two groups.
Fusion between amoebae and swarm cells of dierent
mating types (there are multiple alleles at the mating-type
locus) results in the formation of a zygote. The zygote
nucleus undergoes nuclear division without subsequent cell
division, forming a large, multinucleated cell called a
plasmodium. In some strains, plasmodium formation can
occur in the absence of cell fusion, in which case the entire
life cycle will be haploid. Plasmodia of some species can
become very large ( 4 30 cm in diameter; at least one giant
specimen of P. polycephalum had a surface area of 5.54 m2
(Hausmann and Stiemerling, 1997) and brightly pigmented
and are composed of an interconnected series of tubes
through which cytoplasm ows reversibly at one of the
highest rates of cytoplasmic streaming known in nature
(Figure 1). Fruiting body formation occurs in response to

Slime Moulds

processes, including nonmuscle actin/myosin, microtubules, cytoplasmic streaming, cell dierentiation, cell
compatibility and ageing, and mechanisms of mitosis. All
of the nuclei in a coenocytic plasmodium divide synchronously. The ability to generate, in the laboratory, very large
plasmodia with millions of synchronously dividing nuclei
has made P. polycephalum a valuable organism in the study
of the processes and control of nuclear division (Burland
et al., 1993). P. polycephalum has a haploid chromosome
number of approximately 40 and a genome size of 2.7  108
base pairs (Burland et al., 1993).
Dictyostelid cellular slime moulds

Figure 1 Plasmodium of Physarum polycephalum. The plasmodium has

emerged from a sclerotium, a resting stage that forms when plasmodia are
dried. Reversible cytoplasmic streaming occurs in the tubular structures.
Magnification  4. Photomacrograph & R. L. Blanton.

light and changed environmental conditions. Meiosis

occurs after spore formation, returning the organism to
the haploid state. The most extensively studied species of
plasmodial slime mould is P. polycephalum. Others of note
include Fuligo septica, which forms notably large plasmodia and fruiting bodies and is commonly encountered in
gardens; P. cinereum, whose greyish fruiting bodies formed
on a lawn give cause for unnecessary alarm; Stemonitis,
which forms clusters of fruiting bodies resembling tufts of
hair (Figure 2); and Didymium iridis, which has been used
extensively in mating-type studies.
P. polycephalum has been a useful experimental organism in the study of a variety of fundamental cellular

Figure 2 Fruiting bodies or sporocarps of the plasmodial slime mould

Stemonitis. Magnification  4. Photomacrograph & R. L. Blanton.

The dictyostelid cellular slime moulds comprise over 40

species, three genera and two families. The fruiting body of
a dictyostelid cellular slime mould is called a sorocarp (the
spores are not contained within a common wall but are
suspended in a matrix of slime; the mass of spores is called a
sorus). The unaided eye can see the fruiting bodies of most
dicytostelid species in the laboratory culture dish; however,
they are nearly impossible to see in nature, except in rare
situations when the fruiting bodies are formed in concentrated groups.
Spores germinate to produce a single amoeba. In some
species, amoebae can form cysts. The amoebae multiply
and eventually exhaust their bacterial food supply, at
which point they begin to aggregate together. In D.
discoideum, the signalling system responsible for aggregation is based upon cyclic adenosine monophosphate
(cAMP) signals and has been extensively studied (Parent
and Devreotes, 1996). A detailed discussion of the cAMPbased chemotactic system in D. discoideum is beyond the
scope of this article. Aggregation in a population of
starving cells is induced, some cells beginning to secrete
pulses of cAMP. The initiation of this process is still not
well understood. Other cells respond to cAMP by
migrating towards its source and by beginning to secrete
cAMP themselves. In this manner, the original signal is
relayed far beyond the reach of the original signal. Cells are
nonresponsive to cAMP for a period after their response to
it; this prevents cells from backtracking in response to
cAMP relayed by cells further away from the aggregation
centre. The cells also have membrane-bound and secreted
forms of cAMP-phosphodiesterase, which serves to break
down the cAMP, ensuring that new pulses will be detected
by cells and not obscured by an increasingly higher
concentration of cAMP.
Feeding cells do not stick to one another, but aggregating cells produce cell adhesion molecules that result in endto-end and side-to-side attachment of the aggregating cells,
leading to the streams of cells that distinguish aggregation
in the dictyostelid cellular slime moulds from that in the
acrasid cellular slime moulds.
The aggregating cells form a mound, which elongates to
produce a worm-like structure called a slug, pseudoplas3

Slime Moulds

modium or grex. The multicellular slug is migratory and is

positively phototactic and thermotactic. In nature, this
would result in the slug moving from the humus layer to the
top of the forest litter, thereby maximizing the potential
dispersion of the spores. The slug is surrounded by a slime
sheath, which is left behind as a slime trail as the slug
migrates. Upon reaching the proper environmental conditions, the slug stops migrating and forms a fruiting body
that consists of a delicately tapering stalk and an apical
mass of spores. Two major cell types can be detected in the
slug: prestalk and prespore cells. The prestalk cells
synthesize a stalk that consists of a cylindrical tube of
cellulose and proteins surrounding a population of stalk
cells that are dead at maturity. The stalk tube is synthesized
by the group of prestalk cells as they migrate up the outside
of the tube. The cells at the apex of the growing stalk tube
enter the tube, where they vacuolate, enlarge, form
cellulose walls and die. The combined upwardly directed
force of stalk cell expansion and the creeping of the
prestalk cells along the stalk tube serves to drag the
prespore cells to the apex of the stalk. The prespore cells
synthesize walls containing layers of protein and cellulose
and become spores.
The life cycle just described is asexual and most
commonly involves cells that are haploid. In certain
conditions, including darkness and excess moisture,
aggregating cells will break apart into multicellular clumps
and form thick-walled macrocysts. Fusion between two
amoebae occurs inside the developing macrocyst and the
resulting zygote ingests the other amoebae inside the
macrocyst. The resulting giant cell then undergoes meiosis
followed by multiple mitoses, so that a germinating
macrocyst will yield a number of haploid amoebae.
D. discoideum is the most studied of any slime mould. It
is an important research organism in the study of a variety
of fundamental cellular processes, including cell motility,
cell dierentiation, cell communication and cell signalling
pathways. Among the reasons for the success of D.
discoideum as a research organism are:
. it is easy to grow and manipulate in the laboratory;
. it is haploid, which permits the direct selection of
mutants;
. growth is completely uncoupled from development (i.e.
cell division does not occur during development);
. developmentally lethal mutants can be maintained as
amoebae because development is an induced event;
. development proceeds synchronously as a series of welldened stages over the course of 24 h (Figure 3);
. a physical map of the genome is available, a cDNA
sequencing project is in progress, and the complete
genome sequence is nearing completion;
. a number of molecular genetic tools can be used,
including DNA-mediated transformation, targeted gene
disruption, and tagged random insertional mutagenesis
(Kuspa and Loomis, 1994).
4

Figure 3 Scanning electron micrograph of various developmental stages

of Dictyostelium discoideum, beginning with the flattened consolidated
aggregate in the front right and proceeding clockwise to the mature
fruiting body at the far right. The slug is offset to the front left, indicating
that slug formation is nonobligate. Magnification  40. Photomicrograph
& M. J. Grimson and R. L. Blanton.

Unfortunately, the macrocysts formed by D. discoideum

amoebae are dicult to germinate, meaning that sexual
genetics is not currently possible in this organism. D.
discoideum has a haploid chromosome number of six with a
genome size of 3.4  107 base pairs. It is estimated to have a
total of 8000 genes, of which 500 are estimated to be specic
to development (Loomis, 1990).
Other well-known dictyostelid cellular slime moulds
include Polysphondylium, which has fruiting bodies with
multiple whorls of sori spaced at regular intervals along the
stalk, and D. mucoroides, the most commonly isolated
dictyostelid, with a slug that forms a stalk as it migrates.
The genus Acytostelium is noteworthy because it produces
fruiting bodies with an acellular stalk, the spore mass being
supported only by a cellulosic stalk tube.

Acrasid cellular slime moulds

The acrasid cellular slime moulds number 16 species in six
genera and four families. Based upon dierences in
amoebal types, fruiting body forms, ultrastructural characteristics and type of mitochondrial cristae, the four
families are unrelated and would more appropriately be
classied with various groups of nonfruiting amoebae.
This step has already been taken with the family Acrasidae
(containing the genera Acrasis and Pocheina), which was
combined with the vahlkamid amoeboagellates to create
the Heterolobosea (Page and Blanton, 1985).
The acrasid cellular slime moulds share a similar life
cycle pattern. The trophic phase of each is an amoeba and
in most species the amoebae can form cysts. In three of the
families (the Acrasidae, Guttulinopsidae and Copromyx-

Slime Moulds

idae) the amoebae are said to have a limax form

(cylindrical, not attened). Their pseudopodia lack subpseudopodia and their movement is rapid and smooth. In
contrast, the eumycetozoan amoeba is attened, its
pseudopodia have ne subpseudopodia, and its movement
is slower and more erratic. A fourth family often
considered with the acrasids is the Fonticulidae, as
represented by Fonticula alba. Fonticula amoebae dier
from other acrasid amoebae by being attened and having
subpseudopodia; however, they dier from eumycetozoan
amoebae by having plate-like cristae in their mitochondria,
which is also the case of the mitochondria of the Acrasidae
and Guttulinopsidae. The Copromyxidae have tubular
mitochondrial cristae.
Aggregation by acrasid cellular slime moulds is nothing
like that in the dictyostelids: there are no cell streams, no
cell stickiness and no cell elongation or orientation.
Acrasid aggregation involves the movement together of
cells individually or in small groups. Sorocarp formation is
also distinct from that found in the dictyostelids. A stalk
tube is not formed, there is no division of labour between
cells involved in forming the sorocarp, spores and stalk
cells are not highly dierentiated (they are indistinguishable in most species), and the stalk cells are viable in most
species.
The best known species of acrasid cellular slime mould is
Acrasis rosea. It forms pink sorocarps with branched
uniseriate chains of spores that are distinguishable from
the stalk cells by the presence of distinctive structures,
called hila, at the contact points between spores. The
related Pocheina rosea is notable for its production of
agellated cells.

Phylogenetic and Evolutionary

Considerations
Phylogenetic relationships of the slime
moulds
There are three major phylogenetic questions concerning
the slime moulds. (1) Which of the organisms that we call
slime moulds are related closely enough to be classied
together? (2) Which of the numerous nonfruiting amoebae
groups contain the relatives of the various groups of
fruiting amoebae? (3) Where are the slime mould-containing groups located in relation to the divergence points of
animals, plants and fungi? Answering these questions is
hampered by a lack of data. Detailed ultrastructural
studies have been illuminating, but are time-consuming
and have been accomplished with relatively few species in
the detail that is useful phylogenetically. Sequence data
have been obtained from only a few species, making
determinations of relationships dicult. For example,
sequence analysis of the small subunit ribosomal deoxyr-

ibonucleic acid (rDNA) from plasmodial and dictyostelid

cellular slime mould representatives did not support their
being related. With the subsequent addition of sequence
data from a protostelid, the other slime moulds clustered
together (Spiegel et al., 1995).
Ultrastructural characters have proven to be useful in
phylogenetic studies of a variety of organisms. These
characters include type of mitochondrial cristae, presence
or absence of Golgi bodies, distinct relationships between
mitochondria and rough endoplasmic reticulum, and
whole sets of characters derived from detailed studies of
the mitotic and agellar apparatuses. Detailed study of
trophic stages in the light microscope has also yielded
useful characters, including amoebal form, type of
amoebal locomotion and pseudopodial type. These types
of data provided strong evidence for the relatedness of the
agellated protostelids and the plasmodial slime moulds
and for the creation of the Heterolobosea, as discussed
above.
The Heterolobosea has been conrmed by phylogenetic
analysis of gene sequences for an essential enzyme,
glyceraldehyde-3-phosphate dehydrogenase (Roger et al.,
1996). This study also conrmed the unrelatedness of the
dictyostelid and acrasid cellular slime moulds (Roger et al.,
1996). The Eumycetozoa has been conrmed as a valid
grouping by comparisons of sequences of the protein
synthesis elongation factor 1a (EF-1a) (Baldauf and
Doolittle, 1997). There are insucient data to identify
with condence nonfruiting amoebal or agellate relatives
for the Eumycetozoa or for the acrasid cellular slime
moulds not included in the Heterolobosea.
The consensus from a variety of sequence analyses
indicates that the Eumycetozoa are more closely related to
the fungi and animals than to the green plants (Baldauf and
Doolittle, 1997; Baldauf, 1999). There are conicting
results concerning the location of the branch points for
the three groups, with some data suggesting that the
Eumycetozoa are an outgroup to the animalfungal clade
and other data suggesting that the fungi are the outgroup
of an animaleumycetozoan clade (Baldauf and Doolittle,
1997; Baldauf, 1999). The Heterolobosea may represent
one of the earliest branch points on the eukaryotic lineage
(Cavalier-Smith, 1993).

Evolutionary considerations
Slime moulds are fascinating and worthy of study for their
own sake; however, they would be little more than
curiosities of nature to the broader scientic community
if D. discoideum had not proven to be so amenable to
laboratory study and molecular genetic manipulation.
Modern slime moulds are probably the descendants of the
rst eukaryotic organisms to evolve multicellular development. Many of the processes we associate with multicellularity cell movements during development, cell
5

Slime Moulds

dierentiation, cell-to-cell signalling, formation of cellcell

junctions, cellextracellular matrix interactions are
demonstrated by D. discoideum. It is remarkable that some
of these processes, and the proteins involved in them, are
conserved between D. discoideum and multicellular animals, even such higher organism characteristics as
phosphotyrosine-based signalling (Darnell, 1997) and
cellcell junctions (Grimson et al., 2000). At the same
time, D. discoideum has some profound dierences from
higher multicellular organisms. For example, all cells in a
multicellular animal arise from a single cell, the zygote. The
multicellular stage of dictyostelid cellular slime moulds
arises by aggregation of cells these cells can all be
identical, resulting from divisions of a single progenitor
cell, or they can be from dierent subpopulations of cells, a
phenomenon that has interesting implications for competition between cell lineages and the evolution of development.
The diversity of organisms showing the slime mould
habit presents a fascinating systematic challenge. Understanding the relationships within the fruiting groups and
between the fruiting organisms and their nonfruiting
relatives is important if we are to realize fully the great
potential of these organisms to provide insights into the
evolution of multicellularity and developmental mechanisms.

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Further Reading
Bonner JT (1967) The Cellular Slime Molds. Princeton, NJ: Princeton
University Press.
Buss LW (1987) The Evolution of Individuality. Princeton, NJ: Princeton
University Press.
Kessin RH (2001) Dictyostelium: Evolution, Cell Biology, and the
Development of Multicellularity. Cambridge, UK: Cambridge University Press.
Lee JJ, Hutner SH and Bovee EC (1985) Illustrated Guide to the
Protozoa. Lawrence, KS: Society of Protozoologists.
Loomis WF (1975) Dictyostelium discoideum: A Developmental System.
New York: Academic Press.
Loomis WF (ed.) (1982) The Development of Dictyostelium discoideum.
New York: Academic Press.
Maeda Y, Inouye K and Takeuchi I (eds) (1997) Dictyostelium: A Model
System for Cell and Developmental Biology. Tokyo: Universal
Academy Press.
Margulis L, Corliss JO, Melkonian M and Chapman DJ (eds) (1990)
Handbook of Protoctista. Boston, MA: Jones and Bartlett.
Martin GW and Alexopoulos CJ (1969) The Myxomycetes. Iowa City:
University of Iowa Press.
Olive LS (1975) The Mycetozoans. New York: Academic Press.
Raper KB (1984) The Dictyostelids. Princeton, NJ: Princeton University
Press.