Myxomycota
I. Introduction
These organisms - the slime molds - have been considered as animals,
plants, and as protists. They are probably best regarded as fungi, though
a good case can be made for calling them protists.
II. Labyrinthulomycetes - Net Slime Molds
This is a small, poorly known group. One form, Labyrinthula macrocystis
is thought to cause a serious disease of eelgrass (Zostera marina),
which had a serious impact on brant, redhead, flounders, cod and scallop.
III. Myxomycetes - Plasmodial Slime Molds
A. Habitat
The plasmodial stage is found on rotting
logs, leaf litter, or in moist, shaded soil. The plasmodium may reach several
centimeters in diameter. Some forms are brightly colored, though slime
molds in general are rather inconspicuous members of the woodland scene.
B. Importance
Some slime molds are serious plant pests. One of
the best examples is the cabbage clubroot disease, caused by Plasmodiophora
brassicae, an endophytic parasite. In this disease, zoospores enter
the epidermal cells of the seedling, where they cause abnormal growth forms,
which stunt, and even kill the plant. Of the secondary bacterial infections
can lead to further problems.
On the positive side, slime molds are significant
decomposers, and thus contribute to the maintenance of nutrient cycling.
C. Life cycle of Physarum, a typical plasmodial slime mold.
1. Plasmodium.
The vegetative state is the multinucleate, pigmented,
naked protoplast known as the plasmodium. This yellow protoplast can reach
a size of more than 30 cm.
As it grows over its substrate, it ingests bacteria,
etc., and also absorbs soluble food directly into the mass. It can be seen
to consist of vein-like regions where rapid flow of cytoplasm can be seen.
The flow is erratic, sometimes slowing, stopping, or reversing.
As long as environmental conditions are favorable,
the plasmodium will continue to increase in size, and add nuclei by more
or less simultaneous mitotic divisions of the nuclei.
When temperature or moisture are reduced below acceptable
levels, the plasmodium hardens into a sclerotium. This dormant stage
is composed of irregular hardened masses of macrocysts, each containing
a number of nuclei. The sclerotia are viable for a period of years, and
often serve as the over-wintering stage.
2. Sporulation
When the food supply of a plasmodium is exhausted
or diminished, the slime mold moves to an exposed portion of the substrate,
and begins the formation of sporangia. An additional requirement
for the induction of sporulation is the presence of light - preferably
in the blue/violet portion of the spectrum. In Physarum, the sporangia
are dichotomously lobed irregular structures, but other species have beautifully
formed delicate sporangia. (See Crowder, National Geographic 1926, p 421)
Typically a sporangium consists of an outer wall,
which may be either delicate or firm. It is termed the peridium.
Inside, one often finds a delicate network of hair-like structures known
as the capillitium, among which one finds the spores.
The stipe or stalk of the sporangium arises from
a thin membranous structure known as the hypothallus.
Sporangial initiation is indicated by protoplasmic
aggregation at some of the higher and drier portions of the substrate.
Within this aggregation, very fine vacuoles are formed, within which there
is an accumulation of non-living matter. This form the capillitium. The
nuclei within the sporangium undergo meiosis, with only one meiospore usually
surviving.
3. Swarm cells and amoebae
The meiospores, as they germinate may produce from
one to several naked protoplasts, which may or may not be flagellated.
The habitat is the determiner of this. If germination is in an aquatic
medium, the spore will form flagellated swarm cells. On the other
hand, if the conditions are drier, then ameboid myxamoebae will
ensue. There seems to be no significant difference between these two forms,
other than the presence of flagellae. The flagellated forms seem to all
be biflagellated, with unequal flagellae. They are also capable of ameboid
movement.
Should either of these forms encounter unfavorable
conditions, they can encyst, and later, upon excystment, produce one swarm
cell or myxamoeba, which will feed by means of pseudopodia at the posterior
end, and multiply by mitosis and cytokinesis.
Eventually pairs of cells will undergo synagamy,
and produce a zygote, which, if formed by flagellated cells, will eventually
loosed the flagellae. The zygote is the beginning of a new plasmodium.
(diagram of Physarum life cycle)
IV. Acrasiomycetes - cellular slime molds
A. Habitat
These slime molds differ from the plasmodial slime
molds in that their vegetative phase consists of uninucleate amoebae, which,
because of their size, are rarely observed.
They seem to be most abundant in moist, rich soils,
especially in manured fields. The group is not well known, but is best
exemplified by Dictyostelium.
B. Life cycle of Dictyostelium.
1. Amoebae
These vegetative cells are very similar to the protistan
Amoeba.
They feed in a similar manner, on bacteria and yeasts, They reproduce by
cytokinesis. There is active debate as to whether or not sexual fusion
of amoebae occurs.
2. Aggregation
When a certain density of amoebae has developed,
the cells begin to converge toward centers of aggregation, in response
to secretion by some or all members of a substance termed acrasin.
This substance has been shown to be cyclic AMP. It appears that some cells
act as founder cells, beginning the secretion, then others, as they aggregate,
also secrete the acrasin, enhancing the aggregation process.
3. Pseudoplasmodium
The aggregation of the individual cells results
in the formation of a miniature slug or pseudoplasmodium, whose length
is about 1 mm. This slug continues as a multicellular structure, with the
individual cells retaining their identity, even though intimately associated
together. (They can be separated by shaking the slug in water.) The slug
migrates for a period of hours to days, depending on the moisture in the
environment. Eventually the sorocarp stage is initiated.
4. Sorocarp
When migration ceases, the pseudoplasmodium condenses
somewhat. And begins to form a sorocarp, or spore-producing body.
The sorocarp is composed of two regions - the stalk-like sorophore,
and at the tip, a cluster of spores in a structure termed the sorus.
Each amoeba in the sorus becomes a walled spore. Each spore will eventually
germinate to form a single amoeba.