3 Hyphochytriomycota
MELVIN S. FULLER
CONTENTS
1. lntroduction ........................... .
II. Occurrence and Distribution ............. .
III. Pathogenic States
and Possible Economic Significance ...... .
IV. Ultrastructural Studies ................. .
A. Zoospore Ultrastructure ................ .
B. Mitosis ............................... .
V. Reproduction .......................... .
VI. Development of the Taxonomy
and Current Classification ............... .
VII. Maintenance and Cui ture ............... .
VIII. Conclusions ........................... .
References
1. Introduction
The Hyphochytriomycota are a small group of
zoospore-producing organisms that, although they
live like fungi, i.e., they are nonphotosynthetic
organisms that usually possess cell walls during
their growth phases and acquire nutrients by
absorption, are more closely allied with algal
groups. The diagnostic feature of the group is the
presence of a single, mastigoneme-bearing flagel-
lum that most often is at the anterior end of the
zoospore as it swims. The Hyphochytriomycota
have been classified with the Protista (Whittaker
1969) and Protoctists (Margulis et al. 1990), but
more recent molecular data place them in a clade
with the Oomycetes and heterokont algae. This
clade has been called Kingdom Cromista
(Cavalier-Smith 1993, and Chap. 1, this Vol., and
Kingdom Stramenopila (Alexopoulos et al. 1996);
others simply refer to the group as stramenopiles
(Leipe et al. 1994). Molecular studies (Van der
Auwera et al. 1995) of ribosomal RNA sequences
of Hyphochytrium catenoides and inverted repeat
Department of Botany, University of Georgia, Athens,
Georgia 30601, USA and Darling Marine Center, Univer-
sity of Maine, Walpole, Maine 04573. USA
sequences of mitochondrial DNA in H. catenoides
73 (Mc Nabb et al. 1988) indicate that the
74 Hyphochytriomycota are close relatives of the
Oomycota (see Dick, Chap. 2, this Vol.). Our
75 own molecular studies (S. Lee and M.S. Fuller,
75
75 unpubl.) of the group support inclusion of the
76 Hyphochytriomycota in a clade with the Oomy-
77 cota and heterokont algae, but suggest that the
Hyphochytriomycota are more closely related to
78
78 some groups of the heterokont algae than to the
78 Oomycota.
79 The Hyphochytriomycota have been called
hyphochytrids as opposed to the chytrids (see
Barr, Chap. 5, this Vol.). While the hyphochytrids
look superficially much like the chytrids,
they differ in having zoospores that swim with a
single, anteriorly directed flagellum that has
mastigonemes. Flagellum type is diagnostic for
this very small group with 4 or 5 genera and about
25 species. Although the group is small, it is
exceedingly important to phylogeneticists because
hyphochytriomycete zoospores have a unique
flagellation among stramenopilus fungi. Karling
(1943) used thallus organization to divide the
group into three families: Anisolpidiaceae for
holocarpic forms; Rhizidiomycetaceae for mono-
centric, eucarpic forms; and Hyphochytriaceae for
the polycentric, eucarpic types. Terms such as
those used above for thallus development and
many other terms used in this chapter originated
with pcrsons studying the Chytridiomycota.
Therefore, the definitions given by Barr (Chap. 5,
this Vol.) should be consulted, as the terms will not
be redefined here. Two members of the group,
Hyphochytrium catenoides (Clay et al. 1991) and
Rhizidiomyces apophysatus (Fuller 1960; Fuller
and Barshad 1960) are known to have cell walls
that contain both chitin and cellulose. However,
the composition of the cell walls should not be
used as a diagnostic feature of the phylum,
because no representatives of other species and
genera have been studied relative to cell wall
composition.
The Mycota VII Part A
Systematics and Evolution
McLaughlin/McLaughlin/Lemke (Eds.)
© Springer-Verlag Berlin Heidelberg 2001
74 M.S. Fuller
II. Occurrence and Distribution
Members of the Hyphochytriomycota have been
found in soil and water, includ ing marine habitats,
and appear to have a worldwide distribution, with
most having been found in temperate and tropical
regions (see Sparrow 1960; Karling 1977). The
Hyphochytriomycota are osmotrophic or necro-
trophic and are likely to be found growing as
saprobes when one examines water and soil
samples. Several members of the group may be
biotrophs (Karling 1977; Sparrow 1960) and occur
as parasites of algae (e.g., Canter 1950; Johnson
1957; Karling 1943) and fungi in marine and
terrestrial habitats. No members of the
Hyphochytriomycota are known to grow anaero-
bically. Few isolations of Hyphochytriomycota
have been reported, and the numbers of known
genera and species are so small that it is not pos-
sible to relate habitat, i.e., occurrence mainly in
soil or water, to a taxonomic grouping as Barr
(Chap. 5, this VoI.) has done for chytrids. Soils sus-
pended in water and water-based collection from
aquatic habitats that are baited with, e.g., pollen,
boiled grass blades, and other plant materials, are
more likely to yield chytrids, but hyphochytrids
such as Rhizidiomyces spp. and Hyphochytrium
spp. will be encountered in lower numbers. The
hyphochytrids found in this manner are most
likely growing on dead plant, animal, fungal, or
algal materials. Hyphochytrium infestans was orig-
inally described (Zopf 1884) as a saprobe on dead
ascocarps of Helotium spp. Karling (1967) found
H. oceanum to be widely distributed in soils of the
South Pacific Islands. It should be noted that H.
oceanum is not a marine fungus. Siang (1949)
reported isolating H. catenoides from air samples
collected in Montreal, Canada, While propagules
of this fungus might become airborne, both Barr
(1970) and Fuller (1990) doubt that H. catenoides
is adapted to air dispersal or that it could survive
for extended periods where water was limiting.
However, since Ayers and Lumsden (1977)
showed that H. catenoides retained its viability in
air-dried soil for over 1 year, it is very probable
that wind erosion resulted in propagules of this
fungus becoming airborne.
Very few hyphochytrids have been reported
from the marine environment. Karling (1943) set
up the genus Anisolpidium for a new marine
fungus with anteriorly uniflagellate zoospores. His
fungus, named A. ectocarpii in recognition of the
host, had an intramatrical thallus that was mono-
centric and holocarpic. Except for the zoospores,
it had the appearance of an Olpidium species.
Anisolpidium ectocarpii parasitizes the vegetative
cells and plurilocular sporangia of the brown algae
Ectocarpus mitchellae and E. siliculosus. Johnson
(1957) collected A. ectocarpii from nearly the
same locale as Karling in Beaufort, North
Caro lina and recorded the water temperature as
17.6 ac and the salinity as 22.9%0. Two other fungi
with anteriorly uniflagellate spores, previously
allied with the Chytridiomycota and Oomycota,
respectively, were placed (Karling 1943) in the
genus Anisolpidium and named A. sphacellarum
and A. rosenvingii. Another genus described
(Artemchuk 1972) from a marine habitat is Elina,
that grows epibiotically on gymnosperm pollen.
Members of the genus Elina were characterized
as having monocentric, eucarpic sporangia.
Artemtchuk (1972) recognized his genus as dis-
tinct, because the spores cleaved in the sporangia
and were released, unlike any other member of
the group, by deliquescence of the whole zoospo-
rangial wall. Two species, E. marisalba and E.
sinorifica, were recognized on the basis of
zoospore and thallus sizes. Karling (1979), who
opposed use of the site of zoospore cleavage as a
generic characteristic, concluded that Elina was
not sufficiently different from Rhizidiomyces to
warrant a new genus, and abolished Elina. It the
fungus is rediscovered and spore rele ase is by del-
iquescence of the original sporangial wall, 1 would
favor retaining the genus. Until someone else has
an opportunity to study the fungi seen by Artem-
chuk (1972), 1 favor designat ing Elina as a ques-
tionable genus. Artemchuk and Zelezinskaya
(1969) described another marine hyphochytri-
omycete, Hyphochytrium peniliae, occurring at
epidemic levels on marine crawfish larvae in the
White Sea of Russia.
Although Rhizidiomyces is often isolated
from soil, 1 have found it in water collections and
isolated it twice from freshwater aquaria. Cer-
tainly some of the hyphochytrids are common
inhabitants of the soil and may even serve to
control populations of other soil fungi, namely
Oomycota (Dick, Chap. 2, this VoI.) and Zygomy-
cota (Benny et al., Chap. 6, this Vol.). The occur-
rence of Rhizidiomyces may be controlled by
mycoviruses. Dawe and Kuhn (1983a) have shown
that R. apophysatus contains virus like particles.
The virus can be seen in the nuclei of cells that are
thermally shocked, nutritionally stressed, or aged.
Under these conditions, 50% (Dawe and Kuhn
1983a) of Rhizidiomyces cells can be destroyed.