Morphological observations are given for Colpodella pugnax Cienk., Protomonas amyli Cienk., Protomonas spirogyrae Borzi and Protomonas huxleyi Haeckel

Zopf, W.

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Freshwater Biological Association 
FBA Translation (New Series) No. 151 
Title: Colpodella pugnax CIENK, Protomonas amyli CIENK, Protomonas spirogyrae BORZI 
and Protomonas huxleyi' HAECKEL. 
Author(s) ZOPF W. 
Reference: in: Die Pilzthiere oder Schleimpilze pp 115-124. Breslau. Eduard Trewend 
Original language: German 
of publication of original: 1885 
Translator: J.E. Wightman 
Date of publication of translation: 1981 
Number of pages translation: 5pp 
FBA Translations are to be considered as "provisional". As a ruin 
they have not been prepared by expert translators, nor have, they 
been edited by the original authors. 
They are available from The Librarian, 
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FBA TRANSLATION (NEW SERIES) No. 151 
ZOPF, W. (1885) 
Colpodella pugnax CIENK, Protomonas amyli CIENK, Protomorias spirogyrae BORZI 
and Protomonas huxleyi HAECKEL. 
in: Die Pilzthiere oder Schleimpilze 
pp 115-124. Breslau. Eduard Trewendt. 
Translated by: J.E. Wightman. 
II Monadineae zoosporeae CIENK 
Of the development stages we find here:- 1. the zoospore stage, the 
chief characteristic of the complete group, 2. the amoeba stage, 3. the 
Plasmodium stage, 4. the fructification in zoospore paired zoocysts or in 
swarm-soris, 5. the fructification in spore cysts or in naked spores. 
Family I. Pseudospores 
Out of the following family, the Gymnococcaceae, the pseudospores only 
deviate essentially in the point that the resting spores exist in cysts. All 
pseudospores feed on the contents of green algae or diatoms into whose cells 
they creep. At the fructification, the ingesta is not expelled as a rule. 
Genus I. Colpodella CIENKOWSKI 
The development cycle comprises:- 1. the swarm or zoospore form, 2. the 
zoo-cyst form, 3. the resting spore-cysts. The amoeba stage appears to be 
absent. The way in which the zoospores are liberated is characteristic: the 
tender inner skin of the zoocyst turns itself inside out like a sack and the 
zoospores drift into it, in order to bore through it later. The resting 
spores exist within a cyst. 
Colpodella - pugnax (after CIENK & Nature) a. cell of Chlamydomonas pulvisculus, 
with three zoospores attached, b. Chlamydomonas cell with a zoospore about to 
take nourishment, c. the green content of the algae begins to enter the body 
of the zoospore, d. it is nearly all transferred, e. f. the zoospore has 
transformed to a spherical zoocyst. Inside one can see the already brown 
chlorophyll ball, g. differentiation of the cyst content into zoospores, 
h. i. liberation of the zoospores, k. resting spore developing cysts. 
(All figures by CIENKOWSKI are greatly enlarged) 
2 
1 1. Colpodella pugnax: CIENK 
According to CIENKOWSKI, the zoospores possess a curved sickle shaped 
body, tapered at both ends, a contractile vacuole, and a terminal lash in 
the, vicinity of which a distinct cell nucleus lies. It attaches itself by 
the end remote from the cilium against algae cells (Chlamydomonas pulvisculus), 
pierces through the cellulose membrane of these, and takes the whole 
chlorophyll content of the algae into its thereby distended body. Then it 
releases the emptied cell to drift away. Its form is now that of a Colpoda 
(Infusoria type) with tapered tip which it carries in front of it during the 
swarm. Often several Colpodella zoospores organize a plundering venture. 
Some time after the taking of food, which is agglomorated into a round ball, 
the zoospore become passive, rolled up into atmosphere which surrounds itself 
with a firm membrane, while the foodball, surrounded by a vacuole, takeSon a 
red-brown colour. In the meantime, its plasma fissures into zoospores, the 
sphere becomes a zoocyst. However the emergence of the zoospores is carried 
out in a different way than that of Pseudospora types. That is to say the 
cyst skin bursts and turns inside out, presenting a tender inner skin into 
which the zoospores drift. After gelatination of this, the zoospores are set 
free. Apart from the zoocyst, one distinguishes the resting-spore producing 
cysts. They develop, one to each zoospores takes on spherical form and 
surrounds itself with a membrane. Inside it, the plasma contracts to a 
spherical or ellipsoidal resting spore, next to which the foodball is always 
missing. The germination has not yet been observed. 
(Genus 2 Pseudospora CIENK was not required in this translation - pp 117-120 
JW. Trans). 
Genus 3. - Protomonas 
Essentially only different from the foregoing genera, by the development 
of the fusion plasmodia. 
1. Protomonas amyli CIENK 
It has its home in stagnant freshwater (ponds, swamps, lakes) and must 
be a frequent phenomenon in these places. By an indirect method it is simple 
to confirm this statement. From any particular locality, allow any algae 
(cladophorae, vaucheria, spirogyrae, oedogoniae, charae, nitellae etc.) to 
decompose under water for some time and then add to the infusion pieces of 
starch rich plants like fresh potato tubers, beans, wheat etc. Then one 
finds, after one or two weeks, the organism in this substrate, where it begins 
to consume the amylum grains. Starch seems to be its main and preferred food, 
as all free amylum grains in each infusion used were quickly attacked.2 
From this it follows at the same time, that the Protomonas can exist as a 
saprophyte as well as a parasite. 
The development up to the germination of the resting spores is clearly 
interpreted by the valuable investigations of CIENKOWSKI. 
In order for the zoospore to achieve the liberation, it is provided with 
1 "Contributions to the knowledge of Monads", (Max Schultze's Archive 1), 
2 Ref. the cultivation, it should be noted that it is advisable to introduce 
the starchy substrate in small quantities to the culture, otherwise the 
development of the Protomonas to a rich fission fungus will as a result be 
easily hindered or suppressed. The peelings of fresh potatoes give the best 
substrate). 
3 
two cilia which are so inserted that either one of each lies at each pole, or 
both are, situated together. At the stage of its most active swarm movement, 
the plasma body looks strongly stretched out either spindle formed or worm 
shaped, or otherwise of very variable, often irregular form. 
The Protomonas' takes up nourishment already in the zoospore condition 
and indeed it seeks out the smallest starch particles, for its plasma body 
dimension is as yet too limited to master the large grains (fig 46.2). When 
the zoospore has thrived to two or more times the original size, it transforms 
to the amoeba stage, in which it is able to take up large grains or quantities 
of small particles (fig 46, 3.4). Occasionally this ingestion is rejected 
again, the plasma appears then quite transparent and allows the nucleus to 
be seen clearly, as well as several vacuoles which had earlier been obscured 
by the starch. Ingestion-free younger amoebae usually send out long thin 
pseudopodia (actinophrysform). As they get older and laden with food, the 
pseudopodia development is not so obvious any more. Only relatively short 
pseudopodia are sent out from the surface occasionally; in the end they 
diminish to barely undulating, and eventually complete rounding and skin 
formation occurs (fig 46.5). The previously vacuolish plasma collects into 
a thick grained peripheral layer (v) and fissures into zoospores (vi). 
Thus arises a zoocyst out of a zoospore. The sizes and shapes of the 
zoocysts are extraordinarily varied; alongside spherical, ellipsoidal and 
pear shaped forms (fig 46.1 a,b) one finds laced, club shaped, flask shaped, 
elongated tube shaped and often quite irregular forms, even if these do not 
come from the same culture. 
Protomonas amyli CIENK (mag. approx. 600) 
1. A cell from a potato tuber attacked 
by parasites. The cell membrane is 
very swollen. In its interior one can 
see eight zoocysts of various sizes 
and forms which enclose one or more 
starch grains. The zoocysts marked 
'b' and 'c' have already released their 
zoospores, those marked 'a' exhibit a 
plasma wall cover not yet fissured into 
zoospores; 's' isolated starch grains. 
2. Twincilia zoospores, one with one 
very small starch grain and the other 
with two, 3. Amoeba condition with a 
large starch grain the dark spherical 
tiny body is the nucleus. 4. Amoeba 
with two starch grains. 5. Zoocyst 
with the continuous plasma wall not 
yet fissured into zoospores. 6. Zoocyst 
whose plasma is already divided into 
zoospores. 7. Zoocyst already partly 
emptied, with two zoospores, one of 
which is about to be liberated. 
8. Spherical sporocyst and spore, the 
sporocyst with papillary projections. 
There is however yet another mode of formation of the sporangium. As 
CIENKOWSKI points out, several zoospores can fasten themselves onto a large 
starch grain and after they have withdrawn their cilia, - consequently gone 
into the amoeba stage -, they merge onto the surface of the grain to a 
complete unified plasma layer, - a Plasmodium as CIENKOWSKI rightly called 
it. This exhibits a merely tender shell around the starch grain at first, 
which however gradually gains thickness as it feeds from the starch grain 
4 
elements (it often develops lopsided). What is particularly characteristic 
of this Plasmodium is the lack of any pseudopodia development and with it 
the active movement. Later the Plasmodium surrounds itself with a membrane 
and fissures into more or less numerous zoospores which seek to escape in the 
known manner, namely by boring holes in various places in the membrane and 
forcing themselves through them. 
As far as is known, no more, detailed conditions of this interesting case 
exist, than that before the zoospore development occurs the plasma of the 
spore cyst creeps out in the form of a single large amoeba (or Plasmodium) . 
It shows marked pseudopodia development, consequently strong amoeboid movement 
and occasionally a tendency to develop long fine threads, which here and 
there in their progress show spindle shaped or irregular concentrations of 
the plasma. All of these reasons remind one vividly of the Plasmodium of the 
higher mycetozoae. 
Finally, CIENKOWSKI has also found the. resting spores of developed cysts 
which I have also obtained in nearly all of my cultures. Their development 
is exactly the same as that of the zoospore at first, but eventually the 
plasma which would otherwise have been used for zoospore development, collects 
itself into a spherical or elongated body surrounded by a thick membrane in 
which reserve plasma in the form of compactly accumulated grains lies stored. 
Also the cyst wall thickness becomes papillated with mostly ragged uneven 
projections on the inner surface (see fig. 46.8 and fig 11.2 (see last page)) 
and finally turns brown. Sometimes expelled starch excess lies between the 
cyst membrane and the resting spore. It remains to be discovered how the 
spores behave at the germination. 
That the Protomonas amyli really feeds on ingested starch demonstrates 
that most universally strong corrosion acts on the starch grains which can 
lead eventually to the disappearance of the grain. However in most cases 
only the peripheral layer of the grain is dissolved, obviously as a result of 
the separation of an enzyme. 
2. Protomonas spirogyrae BORZI 1 
According to its complete course of development, as well as the character 
of its individual phases and finally, regarding its biological behaviour, this 
is a typical pseudospore which fits closely next to the Protomonas amyli. 
Protomonas (Protochytrium) spirogyrae BORZI 
a. zoospore, b. amoeba condition, 
c. two amoebae in the act of fusion, 
d. three amoebae in fusion, e. a plasmodium 
formed from four amoebae; each of the 
four vacuole (v), f. plasmoda or larger 
amoeba with its foodballs established 
out of browned chlorophyll (n) these are 
still green, the plasma not yet broken 
up into zoospores, g. a somewhat further 
developed zoocyst, the foodballs (n) 
pushed to one side, the plasma changed 
into a few zoospores, i. sporocyst. 
(s) resting spore with a central drop 
of reserve plasma, (n) food residue. 
All figures by BORZI. (mag. 350). 
1 (ref;. Nuov. Giron. bot. italiano vol. xvi 1884 NO. 1. p. 5) . 
5 
It lives, like the Pseudospora parasitica CIENK, from the contents of 
certain zygnemae (e.g. Spyrogyra crassa and Zygnema cruciatum). The small, 
single cilium pearshaped zoospores (fig 47A) change into amoeba form (fig 47B) 
which differ from Protomonas amyli in that, according to BORZI, they show only 
weak development of mostly blunt pseudopodia, that is, not an actmophrys form. 
By the merging of two or more amoebae (fig 47 C,D) excellent Plasmodia are 
developed (fig 47E) owing to the lack of pseudopodia. Indeed the amoebae can 
also reach the size of plasmodia without coalescence, merely by taking 
nourishment. Amoebae, like plasmodia feed on chlorophyll and starch of host 
plants and convert the chlorophyll bulk into brownish balls which are either 
rejected during fructification time or remain embedded in the plasma body 
(fig 47.G n) . After the plasma has rounded itself and surrounded itself with 
a delicate membrane, it contracts to one side, the food balls being crowded 
to the other side. Then the plasma fissures into two to twenty zoospores. 
So begins the zoospore developed zoocyst (fig"47.H) whose zoospores (according 
to BORZI) become free from the cyst skin by gelatination. 
BORZI also discovered the sporocyst form (fig 47.I). It conforms with 
those of Pseudospora parasitica and Protomonas amyli in as much as within the 
simple, 20-30 u wide cyst skin, the plasma contracts to a spherical or . 
ellipsoidal smooth 15-20 |i thick spore. The latter displays within it a large 
ellipsoidal body (reserve plasma). The spore cyst skin, in contrast to the 
Protomonas amyli, lacks a localized thickening. The germination method of 
the resting spore observed by BORZI has been described and illustrated already 
on- p. 57. The penetration of the host plant by the zoospores takes place in 
such a way that after the loss of the cilia, it nestles against the membrane 
of the host cell, disolves it on a small transfer position and glides in 
through the resulting opening to the inside. 
3. Protomonas huxleyi HAECKEL' 
(ref. Biologische Studien : book 1. p 169 table vi. figs. 5-8) 
This was discovered by HAECKEL on pelagic diatoms (Rhizosolenia) of the North 
Sea by Bergen. Their biology is as yet unexamined. Also the morphological 
condition needs more exact investigation. According to HAECKEL'S work (in part 
indeed yet to be checked), Protomonas huxleyi develops spherical zoocysts 
(about 0.03 mm dia.). Their contents, consisting of fine grained colourless 
plasma, breaks up into a large number of small pieces (measuring about 0.008 u. 
- 8 u) which change to single cilium, pearshaped zoospores. It is not yet 
known in what manner these latter are liberated from out of the fairly coarse 
cyst skin. Cultivated in water drops on a slide, they change into the 
amoeboid condition, in which they probably divide and eventually take on an 
actinophrys type aspect. Investigation is as yet lacking over the possible 
Plasmodia development as well as over the resting spore development. 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
Notice 
 
 
Please note that these translations were produced to assist the scientific staff of the 
FBA (Freshwater Biological Association) in their research. These translations were 
done by scientific staff with relevant language skills and not by professional 
translators. 
 
 
 


Colpodella pugnax Cienk., Protomonas amyli Cienk., Protomonas spirogyrae Borzi and Protomonas huxleyi Haeckel [Translation from: Die Pilzthiere oder Schleimpilze (ed. W. Zopf), pp. 115-124. Breslau, Eduard Trewendt, 1885]

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Colpodella pugnax Cienk., Protomonas amyli Cienk., Protomonas spirogyrae Borzi and Protomonas huxleyi Haeckel [Translation from: Die Pilzthiere oder Schleimpilze (ed. W. Zopf), pp. 115-124. Breslau, Eduard Trewendt, 1885]

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