Spironema
Spironema Klebs, 1893 (ref. ID; 5727)
[ref. ID; 5727]
Diagnosis; Small to medium sized (about 15-50 um) Spironemidae with kineties in indistinct furrows and terminating near or above mid-body. Posterior end with distinct tail. Cortex soft and slightly plicate; cells highly contractile, showing euglenoid movement. (ref. ID; 5727)
Nomenclature; The paper by Klebs was probably available one year earlier than indicated on the title page of the journal, because both Goodey (1916) and Skuja (1956) cited it as 1892. In the absence of further information we recognize 1893 as the year of publication. (ref. ID; 5727)
Remarks; Differs from Stereonema and Hemimastix mainly by its pronounced contractility. Such a difference is widely accepted to be of generic significance (e.g. Euglena and Phacus Leedale, 1967). It is not clear from Klebs' description of S. multiciliatum whether the kineties extend to mid-body or base of tail. Considering the two other species assigned by us to this genus, it is reasonable to assume that the kineties of S. multiciliatum terminate near mid-body. (ref. ID; 5727)
Type species; Spironema multiciliatum Klebs, 1893 (ref. ID; 5727)
- Spironema goodeyi Foissner & Foissner, 1993 (ref. ID; 5727 original paper)
- Spironema multiciliatum Klebs, 1892 (ref. ID; 4921) or 1893 (ref. ID; 5727) reported author and year? (ref. ID; 2659)
- Spironema terricola Foissner & Foissner, 1993 (ref. ID; 5727 original paper)
Spironema goodeyi Foissner & Foissner, 1993 (ref. ID; 5727 original paper)
Diagnosis
In vivo 20-50x4 um. Body vermiform with short tail about 1/4 of cell length. Capitulum conspicuous. Kineties in anterior 1/8 of cell, each kinety with 3-9 flagella. Nucleus distinctly elongate. (ref. ID; 5727)
Descriptions
From (Goodey, 1916): This highly interisting organism occurred in one culture made from Broadbalk 1865 soil. It appeared both on the surface and at the bottom of the culture. My attention was first attracted to it by reason of its great length and its peculiar method of locomotion. It moved slowly in a very hesitating jerky manner for the most part, but would suddenly exhibit rapid and violent spiral twists commencing at its anterior end and travelling down the body, at which times it was propelled at a reasonably fast pace. It was obvious that the organellae causing the slow jerky motion were situated at or towards the anterior end, though they could not be distinguished under a low power microscope. Towards the posterior end a contractile vacuole could be seen in diastole and systole.
I was able to obtain film preparations which, when fixed and stained, revealed the structure of the organism very clearly. The body is extremely long in comparison with the width, and is dorso-ventrally flattened. It measures anything from 20-50 um in length, and averages about 4 um in width. The middle region is generally the widest part of the body. The anterior end is either rounded or has a lateral knob-like portion on either side. The posterior end is drawn out into a long and exceedingly fine tapering tail, and the contractile vacuole occurs just where the body begins to narrow down. The flagella are numerous and comparatively short. They vary in number from seven to eighteen, and the small the organism the fewer the flagella. They are situated in most cases in two lateral rows towards the anterior end of the body, one row being dorsally and the other ventrally placed. I have carefully noted the disposition and number of the flagella, and find that they are not equally distributed on either side, but exhibit a considerable amount of variation in this respect. Klebs, on his P1.xvi, fig.9c, shows a row of flagella extending backwards on one side as far as the beginning of the tail. I have not found anything like this in my organisms. A few of the organisms were fixed just as they were twisting spirally, and one of these is shown in fig. 48. It will be seen from this that the edges of the body are curved, and that the flagella have their origin close to the edges. Each flagellum arises from a small basal granule or blepharoplast distinctly seen in the stained material. I cannot say whether there is a mouth, and although I watched the creatures in life for a long time, I never saw them take in food. There are numerous large granular bodies, however, in the cytoplasm in many of the forms which appear to be ingested bacteria, and because of this, I am of the opinion that a mouth is present. I believe it is situated towards the anterior end, for I have made out, in some cases, a somewhat lighter area here which might be considered as the mouth. In the greater part of the body the cytoplasm is very finely granular and evenly distributed, but towards the posterior end, in the region of the contractile vacuole, it is frequently much vacuolated. The nucleus is a very interesting structure. It is, in most cases, of considerable length, and is situated about half-way down the body. It consists of a long narrow rod of granular material, frequently one-quarter to one-third of the body in length. Towards the middle of it is placed a circular karyosome of deeply staining chromatin. The extra-karyosomic portion of the nucleus appears to be very little different from the general cytoplasm in staining reactions, and is separated from the latter on all sides by a very narrow clear space. There does not appear to be any nuclear membrane. At all events, I have not made out anything comparable with nuclear membrane of other flagellates and amoebae. From the appearance of the stained examples, it seems that all the chromatin is concentrated in the deeply staining karyosome, there being only small scattered granules in the rest of the nucleus. (ref. ID; 5727)
Remarks
Goodey identified this organism as S. multiciliatum Klebs. However, differences in size and shape (very elongate, conspicuous capitulum) and in biotope (soil) warrant species status. Spironema goodeyi is very similar to S. terricola, but the shape of the nucleus is clealy different; since this character is widely used for taxa separation, we recognize the organism as a new species, Spironema terricola probably has a longer tail than S. goodeyi. However, this is not certain, because Goodey's drawings are from fixed specimens whose tails may have been shrunk. Both species differ from S. multiciliatum in body shape and size and in length of kineties, i.e. by the propotion of the flagellated and non-flagellated area. (ref. ID; 5727)
Type location
Agricultural soil from Rothamsted, Broadbalk (England). (ref. ID; 5727)
Occurrence
As yet recorded only from type location. (ref. ID; 5727)
Type material
Goodey made hematoxylin slides but did not mention where they have been deposited. Probably, they are at the University of Birmingham or at the Rothamsted Research Station. (ref. ID; 5727)
Spironema multiciliatum Klebs, 1892 (ref. ID; 4921) or 1893 (ref. ID; 5727) reported author and year? (ref. ID; 2659)
Diagnosis
In vivo 14-18x2-3 um. body lanceolate with tail about 1/3 of cell length. Capitulum inconspicuous. Kineties with 6-13 flagella each. (ref. ID; 5727)
Descriptions
Klebs, 1893 Translated from German: The body shape is highly characteristic because of the long, acute tail, the two spiraled grooves which commence that anterior end and extend up to the tail filament. The ciliature is very unusual. The edge of each spial groove is provided with numerous small cilia which normally do not move uniformly but independently from each other. In the first specimens I observed cilia only in the anterior portion. Recently, I noticed specimens with cilia up to the posterior end or the beginning of the tail, respectively. I may have overlooked them earlier. The animal moves sluggishly back and forth, occasionally settling with the acute tail. The posterior region of the body including the tail is fairly stiff and changes hardly. In contrast, the anterior portion is highly metabolic, contracts and extends again, bends and curves to all sides. Small bodies are propelled towards the grooves by the movement of the cilia and small bacteria (small greenish bodies) probably sink into the cytoplasm beneath the anterior portions of the groove and become enclosed in vacuoles. The contractile vacuole is in the posterior portion where the tail begins. Unfortunately, the reproduction could not be observed. (ref. ID; 5727)
Remarks
Spironema multiciliatum differs from S. goodeyi and S. terricola in size and shape of body in the absence of a conspicuous capitulum, and in habitat. The nucleus has not been described by Klebs. (ref. ID; 5727)
Type location
Ditch water in Germany. Exact location unknown. (ref. ID; 5727)
Occurrence
As yet recorded only from Germany by Klebs (1893) and Zolffel (pers. commun.). (ref. ID; 5727)
Type material
Not available. (ref. ID; 5727)
Spironema terricola Foissner & Foissner, 1993 (ref. ID; 5727 original paper)
Diagnosis
In vivo 30-50x3-4 um. Body vermiform with very thin tail half as long as cell. Capitulum conspicuous. Kineties in anterior 1/8 of cell, each kinety with about 8 flagella. Nucleus roundish. (ref. ID; 5727)
Descriptions
- Light microscopy: Cells often curved or distinctly bent at mid-body or near origin of tail; non-rigid, may extremely deform; anterior more distinctly flattened than in mid-body; capitulum conspicuous in broad axis view; tail unflattened, very thin and fragile, readily shortens under cover glass pressure and probably also during preparation. Nucleus near centre of thickened portion of cell. Contractile vacuole near base of tail, surrounded by bleb-like vesicles. About 30 tiny (0.8x0.3 um), rod-like inclusions in anterior cell portion, corresponding to posterior compartment of extrusomes. Flagella 10 um long, rather rigid, beat asynchronously. Food vacuoles globular, in mid-body. Occasionally a spindle-like inclusion 3x1.5 um in size. Usually slowly gliding on tail or trembling and moderately fast swimming while rotating about longitudinal axis. Anterior body portion revolves in settled specimens which use tail as anchoring device. Swimming and gliding cells show distinct euglenoid movement from time to time, even if seemingly undisturbed. Cell dilations commence near apex and progress towards tail. Contracted cells conspicuously twisted up to beginning of tail. Movement soon ceases in squeezed cells. (ref. ID; 5727)
- Electron microscopy: Cortical plates slightly plicate. Epiplasm consists of heavily staining osmiophilic sheet (outer layer) and thicker, lighter staining inner layer; continuous in capitulum, gradually reduced toward ends of kineties, therefore not evenly broad as in Hemimastix but tapering. Flagella-free portion of cortex without epiplasm i.e. solely supported by microtubules. Cortical microtubules along short axis continuous over capitulum (loop-like), i.e. originate and end in tail. Cortical microtubules along broad axis originate from both sides of capitulum, viz. slightly beneath cell's apex and from cell membrane near flagellar pits; also extend to tail.
Mean diameter of flagella 312 nm (range 250-350 nm, n=5), axonemes 198 nm (175-214 nm, n=5), basal bodies 183 nm (176-190, n=5), membranous sacs 61 nm (60-62 nm, n=6); mean length of basal bodies 236 nm (227-250, n=4). Extrusomes about 4x0.23 um in size, barely recognizable in light microscope, anchored in epiplasm along kineties and in capitulum. Anterior extrusome compartment ("nail") with thin granular centre, tip cup-like, posterior end ("nail head") with three granule-like thickenings. Posterior bottle-like compartment with two bulbous dilations at anterior end and electron lucent zone separating nail head from content of bottle. Food vacuoles contain electron-dense, amorphous masses and membrane residues. Numerous globular vesicles, about 800 nm in diameter, with homogeneous content of medium electron density scattered throughout cytoplasm. Cisterna of rough endoplasmic reticulum conspicuous along nucleus. (ref. ID; 5727)
Etymology
"Terricola" for "living in soil". (ref. ID; 5727)
Type location
Soil from the Grand Canyon, USA (upper entrance to Bright Angel Trail), 36 degrees N, 112 degrees E. (ref. ID; 5727)
Occurrence
As yet found only at type location, viz, in a pine forest at the edge of the Grand Canyon. Spironema terricola occurred in soil of grass patches scattered in a thick (up to 10 cm) layer of litter on red soil. The soil appeared virtually dry when collected and was air-dried for three weeks. It is thus reasonable to assume that S. terricola emerged from resting cysts. (ref. ID; 5727)
Type specimens
One slide of holotype specimens and three slides of paratypes specimens (all protargol-impregnated) have been deposited in the collection of microscope slides of the Oberosterreichische Landesmuseum in Linz, Austria. Accession numbers: 44, 45, 46, 47/1992. (ref. ID; 5727)