Colpoda Morphology
The World of Protozoa, Rotifera, Nematoda and Oligochaeta
- Glossary
- Toxicity test
- Reference
- Journal list
Class Kinetofragminophora: Subclass Vestibulifera: Order Colpodida (ref. ID; 2013)
Suborder Colpodina Foissner, 1978: Family Colpodidae Ehrenberg, 1838 (ref. ID; 7706) [ref. ID; 2013]
Body distinctly reniform in shape, dorso-ventrally flattened. Right body edge strongly convex, left body edge concave often appearing as through a bite had been taken from it. A shallow diagonal somatic groove (not easily visible) originating on the dorsal surface travels round left side to entrance of vestibulum on the flattened ventral surface. Ciliation uniform in longitudinal or oblique orientated grooves. Several notches which denote ciliary grooves often visible on preoral part of left body edge. Caudal cilia may be present on some species. There is a horse-shoe shaped arc of closely-set cilia on the right of the vestibular entrance. Single rounded macronucleus with 1, 2 or 3 micronuclei. Single terminal contractile vacuole. Division takes place in thin-walled cysts, thick-walled protective cysts also formed. Could be easily confused with Wenrichia.
Quote; Colin R. Curds "British and other freshwater ciliated protozoa Part I Ciliophora: Kinetofragminophora" Cambridge University Press, 1982 (ref. ID; 2013) [ref. ID; 4030]
There are 23 species presently assigned to the genus Colpoda, none of which is multimicronucleate. Among the colpoda-shaped ciliates, species of the genus Tillina also have several micronuclei. This latter genus is further distinguished by its large body size (>120 um), an elongate canal-like oral cavity, and by a unique set of ectoplasmic channels which converge at the contractile vacuole. (ref. ID; 4030) [ref. ID; 7205]
Ciliates of the genus Colpoda are widely distributed organisms, commonly found encysted on most types of vegetation and in most soils. Few studies have carefully examined morphometric variation in the genus. Kahl (1931-1935) provided one of the most complete reviews of species in the genus Colpoda. He redescribed 7 species and described 10 new species, primarily basing his distinctions between species on variations in body morphology. Burt (1940) undertook a careful analysis of intraspecific variation in the genus. The major characters separating the species he studied were somatic length, total kinety number, and characteristics of the Feulgen-stained macronucleus. Lynn (1976, 1977) and Novotny et al. (1977) redescribed five common species using silver impregnation techniques and recorded variability within clonal cultures. Recently, Foissner (1980) provided careful drawings and descriptions of 10 species of which 4 are new. (ref. ID; 7205) [ref. ID; 7435]
The molecular diversity within the D2 domain of the 23S ribosomal RNA molecules of Colpoda. (ref. ID; 7435)
- Colpoda acuta Buitkamp (ref. ID; 2365 original paper)
- Colpoda aspera Kahl, 1926 (ref. ID; 1622, 2288, 3593) reported year? (ref. ID; 1618) reported author and year? (ref. ID; 4609, 7118)
- Colpoda brasiliensis Foissner, 2003 (ref. ID; 4915 original paper)
- Colpoda californica (ref. ID; 1622)
- Colpoda cavicola (ref. ID; 1620)
- Colpoda cucullus (ref. ID; 191, 7118, 7205, 7589)
- Colpoda cucullus Ehrenberg (ref. ID; 3491)
- Colpoda cucullus O.F. Muller, 1773 (ref. ID; 4861) or 1786 (ref. ID; 1219, 1622, 1629, 2245, 3593) reported year? (ref. ID; 646, 1618, 3342, 3698, 4842, 5624) or (O.F. Muller, 1773) Gmelin, 1790 (ref. ID; 4609)
Syn; Colpoda lucida Greeff, 1883 (ref. ID; 1622); Kolpoda cucullus O.F. Muller, 1773 (ref. ID; 4609); Tillina flavicans Stokes, 1885 (ref. ID; 1622) - Colpoda discoidea Gellert, 1956 (ref. ID; 4030)
- Colpoda duodenaria Taylor & Furgason, 1938 (ref. ID; 4030) reported year? (ref. ID; 1618) reported author and year? (ref. ID; 191)
- Colpoda ecaudata Liebmann, 1936 (ref. ID; 1629, 2128, 4609)
Syn; Cyclidium ecaudatum Liebmann, 1936 (ref. ID; 4609) - Colpoda elliotti Bradbury & Outka, 1967 (ref. ID; 4030) reported author and year? (ref. ID; 191)
- Colpoda eurystoma Gellert, 1950 (ref. ID; 4030)
- Colpoda fastigata (ref. ID; 1622)
- Colpoda flavicans (Stokes, 1885) Foissner, 1993 (ref. ID; 4861)
- Colpoda formisanoi (ref. ID; 4915)
- Colpoda fragilis Powers, 1933 (ref. ID; 4030)
- Colpoda henneguyi Fabre-Dom, 1888 (ref. ID; 1622) or Fabre-Domergue (ref. ID; 4842) reported author and year? (ref. ID; 7118)
- Colpoda inflata (Stokes, 1884) (ref. ID; 2128) or 1885 (ref. ID; 1622, 1629, 3593, 4522) reported year? (ref. ID; 1618, 2342, 3698, 4842, 4848, 5462), (Stokes, 1884) Kahl, 1931 (ref. ID; 4609, 4861) reported author and year? (ref. ID; 191, 4332, 4851, 7205)
Syn; Colpoda rouxi Kahl, 1926 (ref. ID; 1622); Colpoda steini Roux, 1901 (ref. ID; 1622); Tillinia inflate Stokes, 1884 (ref. ID; 4609) - Colpoda irregularis Kahl, 1931 (ref. ID; 3593) reported author and year? (ref. ID; 1622)
- Colpoda lucida Greeff, 1888 (ref. ID; 4861) reported year? (ref. ID; 2128) reported author and year? (ref. ID; 7118)
- Colpoda magna (Gruber, 1880) Lynn, 1978 (ref. ID; 4609) reported author and year? (ref. ID; 1629, 7118)
Syn; Tillina canalifera Turner, 1937 (ref. ID; 4609); Tillina magna Gruber, 1880 (ref. ID; 4609) - Colpoda maupasi Enriques, 1908 (ref. ID; 1622, 4861) reported year? (ref. ID; 1618) or Enriquez, 1908 (ref. ID; 5918) reported year? (ref. ID; 3342) reported author and year? (ref. ID; 65, 191, 7205)
Syn; Colpoda steini Maupas, 1883 (ref. ID; 1622) - Colpoda ovinucleata Foissner (ref. ID; 3698)
- Colpoda patella Kahl, 1931 (ref. ID; 3593) reported author and year? (ref. ID; 1622)
- Colpoda penardi (ref. ID; 1622)
- Colpoda praestans Penard, 1922 (ref. ID; 1622) reported author and year? (ref. ID; 7205)
Syn; Tillina magna Penard (ref. ID; 1622) - Colpoda quinquecirrata Gellert, 1955 (ref. ID; 4030)
- Colpoda reniformis Kahl, 1931 (ref. ID; 3593) reported author and year? (ref. ID; 1622)
- Colpoda rotunda Foissner (ref. ID; 3698)
- Colpoda simulans (ref. ID; 1622, 7205)
- Colpoda spiralis Novotny, Lynn & Evans, 1977 (ref. ID; 4030 original paper) reported author and year? (ref. ID; 7205)
- Colpoda steini Maupus, 1883 (ref. ID; 1335, 1622, 1629, 1796, 2245, 2342, 3116, 3593, 7533) reported year? (ref. ID; 1219, 1618) reported author and year? (ref. ID; 65, 3370, 7564), steinii Maupas, 1883 (ref. ID; 4609, 4613) reported year? (ref. ID; 3342, 3698, 3744, 4842) reported author and year? (ref. ID; 191, 7205)
Syn; Tillina saprophila Stokes, 1884 (ref. ID; 1622, 4609, 4613) - Colpoda tripartita Kahl, 1931 (ref. ID; 4861) reported author and year? (ref. ID; 1622)
- Colpoda triquetrus O.F. Muller, 1786
See; Chlamydodon triquetrus (O.F. Muller, 1786) Kahl, 1931 - Colpoda variabilis Foissner (ref. ID; 3698)
Colpoda aspera Kahl, 1926 (ref. ID; 1622, 2288, 3593) reported year? (ref. ID; 1618) reported author and year? (ref. ID; 4609, 7118)
Descriptions
Cytostome about one-third from the anterior end; 14-16 meridians; anterior keel with 5 indentations; in fresh water. (ref. ID; 1618)Measurements
12-42 um long. (ref. ID; 1618)Colpoda brasiliensis Foissner, 2003 (ref. ID; 4915 original paper)
Diagnosis
Size about 33x18 um in vivo. Body covered with a mineral envelope and Colpoda maupasi-shaped, that is, reniform with anterior half narrower than posterior. Macronucleus usually in posterior dorsal quadrant of cell, ellipsoidal, with single central nucleolus. On average 12 ciliary rows, distance between first and second row right of oral opening distinctly widened anteriorly; number of preoral kineties thus higher at left than right side. Oral apparatus in second quarter of cell, left polykinetid composed of about 5 kineties becoming slightly longer distally. (ref. ID; 4915)Descriptions
When I first saw this species, it was rather abundant but misidentified as a "small, dirty variety of C. maupasi". Two weeks later, when I recognized it as a distinct species, it was already rare. Thus, the morphometric data are incomplete. Further, the protargol preparations are of only mediocre quality because the mineral envelope impregnates rather intensely and hides the ciliary pattern. Thus, the description should be refined by observations on other populations. Size 30-40x15-25 um in vivo, usually about 33x18 um. Invariably Colpoda maupasi-shaped, that is, reniform with flat oral indentation above mid-body and narrowed anterior half; no diagonal (postoral) groove. Macronucleus in rear half of cell, usually in posterior dorsal quadrant, about 7x5 um and very hyaline in vivo, except for the small central nucleolus, which stains red with methyl green-pyronin and impregnates rather intensely with protargol; micronucleus not unequivocally identified, probably attached to macronucleus and ellipsoidal. Contractile vacuole in centre of rear body end. Cortex flexible, special cortical granules (extrusomes) recognizable neither in vivo nor with methyl green-pyronin stain. Mineral envelope 1-3 um thick, composed of clay particles up to 3 um in sized embedded in a slimy matrix to which environmental organic debris and bacteria may adhere; stains bluish with methyl green-pyronin and provides cells with a rough, dirty surface. Cytoplasm colourless, usually packed with food vacuoles 5-10 um across and with many minute crystals conspicuously sparkling under interference contrast optics, as in many other small colpodids and marynids (Foissner et al. 2002). Feeds on bacteria, whose compact spores become distinct in late food vacuoles. Crawls clumsily on soil particles, but may also rapidly glide on microscope slide. Cilia about 10 um long in vivo, paired, except in rear body portion, where only the dikinetid's posterior basal body is ciliated; no elongated caudal cilia; arranged in an average of 12 rows, more densely ciliated anteriorly than posteriorly. Ciliary rows in typical Colpoda pattern with, however, some specializations; abut preorally, except for three postoral kineties, forming narrow suture extending from anterior pole to summit of oral opening. Right side ciliary rows slightly sigmoidal, distance between first and second row right of oral opening conspicuously widened anteriorly. Left side ciliary rows rather distinctly sigmoidal, more narrowly spaced than right side rows along preoral suture, producing a special pattern in that an additional row is intercalated between each two right side rows. Invariably three postoral kineties, leftmost kinety slightly shortened anteriorly. Oral apparatus small and thus inconspicuous, in flat indentation of second quarter of cell, posterior margin of left ciliary field on average 43% distant from anterior body end. Vestibulum conical, about 6 um wide and deep, vestibular opening and oral ciliary fields from conspicuous, circular pattern; pharyngeal fibres not impregnated. Left oral polykinetid indistinctly cuneate, composed of an average of five slightly convex rows becoming slightly longer distally, cilia of proximal rows about 3 um long, those of distal rows 5 um, do not form a distinct beard as in Colpoda steinii. Right oral polykinetid semicircular, probably composed of three or four rows of cilia projecting about 5 um from vestibular margin. (ref. ID; 4915)[Cysts]: Resting cysts observed only in protargol slides, globular to broadly ellipsoidal, about 24x22 um on average. Wall up to 2.5 um thick and not impregnated, covered by the brownish-impregnated mineral envelope of the trophic cell, providing cysts with a rough, irregular surface. Central nucleolus of macronucleus distinct. (ref. ID; 4915)
Remarks
Colpoda brasiliensis highly resembles small specimens of Colpoda maupasi, one of the most frequent soil ciliates (Foissner 1993; Foissner et al. 2002), but is easily distinguished from that species by the mineral envelope (present vs. absent), the structure of the macronucleus (with vs. without central nucleolus), the arrangement of the preoral ciliary rows, and the number of somatic ciliary rows (about 12 vs. 15-40, usually 20-30) and kineties comprising the left oral polykinetid (about 5 vs. 9-10). Several populations of C. steinii, C. formisanoi, and Dragescozoon terricola also resemble C. brasiliensis, especially in body size and shape, the location and structure of the macronucleus, and the preoral ciliary pattern. However, all C. steinii populations have a highly characteristic spoon-shaped, comparatively large left oral polykinetid, conspicuously different from the rectangular or slightly cuneate, minute polykinetid of C. brasiliensis; all C. formisanoi populations have an acute posterior body end, distinctly different from the broadly rounded end of C. brasiliensis; and Dragescozoon terricola has the macronucleus in the anterior body end and a somewhat different preoral ciliary pattern. Of course, all these species, described in detail in Foissner (1993) and Foissner et al. (2002), lack a mineral envelope. On the other hand, they have conspicuous similarities suggesting a common ancestor and resurrecting the genus Paracolpoda Lynn, 1978. Probably, C. brasiliensis is more closely related to the Colpoda steinii group than to the C. maupasi group, with which it shares mainly body shape. (ref. ID; 4915)Etymology
Named after the country in which discovered. (ref. ID; 4915)Type location
Clayic top soil from a small island in the Amazon River near Manaus, Brazil. 04 degrees S 60 degrees W. (ref. ID; 4915)Type specimens
1 holotype and 1 paratype slide with protargol-impregnated trophic and cystic specimens have been deposited in the Biology Centre of the Museum of Upper Austria in Linz (LI), Austria. Relevant specimens are marked by black ink circles on the cover glass. (ref. ID; 4915)Colpoda cucullus O.F. Muller, 1773 (ref. ID; 4861) or 1786 (ref. ID; 1219, 1622, 1629, 2245, 3593) reported year? (ref. ID; 646, 1618, 3342, 3698, 4842, 5624) or (O.F. Muller, 1773) Gmelin, 1790 (ref. ID; 4609)
Synonym
Colpoda lucida Greeff, 1883 (ref. ID; 1622); Kolpoda cucullus O.F. Muller, 1773 (ref. ID; 4609); Tillina flavicans Stokes, 1885 (ref. ID; 1622)Descriptions
Anterior keel with 8-10 indentations; 29-34 ciliary grooves; cilia mostly paired; macronucleus with a stellate endosome; trichocysts rod-form; usually with abundant food vacuoles; in fresh water with decaying plants. (ref. ID; 1618)Measurements
40-110 um long. (ref. ID; 1618)44-50x22-30 um. (ref. ID; 3342)
Colpoda duodenaria Taylor & Furgason, 1938 (ref. ID; 4030) reported year? (ref. ID; 1618) reported author and year? (ref. ID; 191)
Descriptions
12 longitudinal ciliary rows; 3 postoral rows; 2 long cilia at the posterior end; long cilia project out from the cytostome along its posterior margin, forming a "beard"; a contractile vacuole terminal; macronucleus ovoid, with crescentic micronucleus; division into two to eight individuals in division cyst; but no division in trophozoite stage; bacteria-feeder; fresh water. (ref. ID; 1618)Measurements
20-40 (9-60) um. (ref. ID; 1618)Colpoda inflata (Stokes, 1884) (ref. ID; 2128) or 1885 (ref. ID; 1622, 1629, 3593, 4522) reported year? (ref. ID; 1618, 2342, 3698, 4842, 4848, 5462) or (Stokes, 1884) Kahl, 1931 (ref. ID; 4609, 4861) reported author and year? (ref. ID; 191, 4332, 4851, 7205)
Synonym
Colpoda rouxi Kahl, 1926 (ref. ID; 1622); Colpoda steini Roux, 1901 (ref. ID; 1622); Tillinia inflate Stokes, 1884 (ref. ID; 4609)Descriptions
Anterior keel with 6-8 indentations; number of ciliary grooves (or meridians) 21-24; cilia mostly in pairs; in fresh water among vegetation. (ref. ID; 1618)-
[ref. ID; 4332]
- [Vegetative cell]: The vegetative cells (strain HSS-1) are reniform or ellipsoidal, and measures 56.5 um in length by 39.3 um in width on average. The nuclear apparatus consists of a spherical macronucleus and a lenticular or spherical micronucleus. The somatic infraciliature is made up of 18-24 kineties, which are really dikineties as seen by electron microscopy (Lynn 1976). Each dorsal or lateral kinety has 14-24 pairs of kinetosomes. The oral infraciliature presents an ordered left oral field constituted by 11 to 12 short oblique rows of kinetosomes and a right oral field with numerous kinetosomes more or less anarchically disposed. The vegetative cells (strain HSL-1) are always reniform, have a pronounced indentation at the end of anterior third of body, and average 62.0 um in length by 46.2 um in width. There is an oval macronucleus and a lenticular or spherical micronucleus. The somatic infraciliature is composed of 23-27 dikineties. The oral infraciliature is made up of a left oral field with 13-16 short oblique rows of kinetosomes and disordered right oral field. (ref. ID; 4332)
- [Morphogenesis during cell division]: In Colpoda inflata, as in other species of this genus, division produces a cyst and forms two or four tomites. At first, the vegetative cell becomes spherical and gives rise to a very thin division cyst wall. Simultaneously, the micronucleus separates from the macronucleus and begins to divide. Then, the oral infraciliature is resorbed progressively, first, the left oral field and then the right oral field. Only when the oral infraciliature has almost disappeared does the vegetative cell elongate and the somatic pairs of kinetosomes begin to proliferate longitudinally, according to an antero-posterior gradient and from the ventral to the dorsal side. As a result of this proliferation, the somatic kineties present a bipolar pattern. Meanwhile, the micronucleus mitosis is finishing and the macronucleus, situated in parallel to the antero-posterior axis of the cell, elongates. Then, both cellular poles rotate in the opposite direction. As a result of this rotation, the somatic kineties break in two parts at the equatorial level and an unciliated area appears among them owing to the cellular elongation. Soon after the individualization of the two future tomites, stomatogenesis starts simultaneously in both cells. However, the morphogenetical process is quite different when four tomites are originated into a dividing cyst. In such cysts, the two tomites-I first separate because of the stretching of the cell. Then, one tomite-I moves by the side of the other. Later, each tomite-I undergoes the same cortical and nuclear phenomena as those taking place in a single dividing cell. In these cases, the stomatogenesis begins simultaneously in the four tomites. Under certain environmental conditions, we have observed that some vegetative cells do not form the division cyst wall. In this way, the four tomites arrange in a chain pattern because the posterior tomite-I has not moved after the end of the first division round. In these cellular chains, each of the four future tomites undergoes a stomatogenic process; generally these cells are not viable because of an abnormal distribution of the nuclei. In any case, during stomatogenesis the right oral primordium results from the longitudinal and lateral proliferations of the kinetosomes of the anterior extreme of a single somatic kinety. On the contrary, the left oral primordium originates from the longitudinal proliferation of the anterior extremes of four, five, or six somatic kineties. As a result of these proliferation, each tomite presents a right oral primordium composed of an irregular field with numerous kinetosomes and a left oral primordium made up of 4-6 groups, each with three short rows of kinetosomes. Later, these groups of kinetosomes become closer to constitute the definitive left oral field. In the definite right oral field of each tomite, there is a curved double row of kinetosomes and a disordered field of kinetosomes to its right. After comparing the number of rows of kinetosomes composing the left oral field in a vegetative cell with the number of groups composing the left oral primordium in a tomite, we conclude that some kinetosomes of this left oral primordium are resorbed during the last stage of stomatogenesis. With regard to the macronuclei, the macronucleus of each tomite undergoes a chromatinic extrusion process once all the oral infraciliary structures are formed. The chromatinic mass is always smaller than the macronucleus and stains much more intensely. It becomes compact and rapidly diminished in size until it disappears. This process develops nearly or exactly synchronously in each daughter macronucleus. Finally, cytokinesis ensures and the division cyst wall breaks, releasing two or four tomites. (ref. ID; 4332)
- [Cortical and nuclear phenomena during resting cyst formation]: In addition to the division cysts, the different species of Colpoda can produce resting cysts under certain environmental conditions, such as starvation. In C. inflata, the encystment process lasts around 120-180 h. During this cell differentiation process, some cortical and nuclear phenomena develop during the 6-36 h after encystment induction and therefore, before the definite resting cyst wall formation. As in cell division, the vegetative cell first adopts a spherical shape and the micronucleus moves away from the macronucleus. Then, the oral infraciliature is resorbed. In this case, the left oral field is also the last to disappear. Until this stage, the cortical phenomena during resting cyst formation are the same as those of cell division with the exception of the somatic infraciliature. During resting cyst formation, the somatic kineties do not proliferate longitudinally but the kinetosomes lose their cilia and dispose in a radial way. There are fewer kinetosome pairs in the precystic cell than in the vegetative cell. Therefore, we think that some pairs of kinetosomes are resorbed during encystment. Meanwhile, the macronucleus undergoes a chromatinic extrusion process. In this case, the chromatinic mass is also smaller than the remaining macronucleus. Later, this mass is resorbed. After this process, the micronucleus moves closer to the macronucleus again. In the resting cyst the micronucleus is always lenticular. When we induce resting cyst formation in exponential growth, the dividing cells can also produce resting cysts. In these cases, the two macronuclear masses undergo a chromatinic extrusion process. (ref. ID; 4332)
Measurements
35-90 um long. (ref. ID; 1618)Colpoda maupasi Enriques, 1908 (ref. ID; 1622, 4861) reported year? (ref. ID; 1618) or Enriquez, 1908 (ref. ID; 5918) reported year? (ref. ID; 3342) reported author and year? (ref. ID; 65, 191, 7205)
Synonym
Colpoda steini Maupas, 1883 (ref. ID; 1622)Descriptions
Cytostome about one-fourth from the anterior end; anterior keel with 5 indentations; 16-18 meridians; in fresh water. Bensonhurst strain has been reported to produce several types of cysts, due probably to nutrition, age and size of the trophic forms. (ref. ID; 1618)-
[ref. ID; 5918]
- [Light microscopy]:
- Body shape. Colpoda maupasi appears slightly reniform when viewed laterally. When viewed ventrally and dorsally the ciliate appears pyriform, shaped like a 'teardrop'. The short preoral keel and anteriorly placed oral cavity give this species a distinctively elongate shape. The postoral portion of the body may become swollen with densely packed food vacuoles. (ref. ID; 5918)
- Kinetome and argyrome. C. maupasi is completely holotrichous. There are no somatic regions where kinetosomes are usually absent as described for C. steinii (Lynn 1976). The anterior ends of the left keel kineties terminate on an anterior convergence region which is almost dextral. Several kineties insert on the dorsal portion of the convergence region or keel. The kinetal counts for various body regions are quite constant within this clone. (ref. ID; 5918)
- Oral cavity. The right and left oral polykinetids are situated, respectively, on the right and posterior walls of the oral cavity. They are oriented about 60 degress to each other when viewed ventrally. The coniform oral cavity extends dorsad and slightly to the right. Often it is coursed by fine siver lines. (ref. ID; 5918)
- Expulsion vesicle pore and cytoproct. The expulsion vesicle pore is located in the extreme posterior left sector. Upon it, most of the somatic kineties converge. No cytoproct was observed, although often slightly anterior to the expulsion vesicle pore, denser silver lines were sometimes present. (ref. ID; 5918)
- Nuclei. The spherical macronucleus has a complexly ramified Feulgen-negative 'body'. Feulgen-postivie material is distributed among these ramifications. A single crescentic micronucleus is closely apposed to the macronucleus. (ref. ID; 5918)
- [Electron microscopy]:
- Membrane systems. A unit cell membrane covers the entire organism. It is subtended by alveoli on all surfaces with the exception of ciliary shafts and pits, expulsion vesicle pores, cytopharynx, parasomal sacs, and between the bases of oral polykinetidal cilia. The alveoli are unit-membrane-bound sacs which lie above the epiplasm. Alveolar junctions, formed by the apposition of adjacent alveolar boundaries, are filled with electron-dense material resembling that of the epiplasmic matrix. After fixations using phosphate buffer, the alveoli are indistinct. The epiplasm is a single homogeneously electron-dense layer about 26 nm in thickness, which underlies the plasmalemma or alveoli of all cortical structures with the exception of the cytostome, parasomal sacs, and the bottom of the expulsion vesicle pore. As in other Colpoda species (Lynn 1974), the epiplasm is continuous with the terminal plate of somatic kinetosomes. The cortex is ridged between somatic kineties. Anterior transverse, posterior transverse, and postciliary microtubular ribbons course within a ridge. These microtubules are closely associated with the epiplasm, perhaps by filamentous material. (ref. ID; 5918)
Kinetosome. Kinetosomes are observed as linked pairs. In most sections the cilia are in pairs, although in some organisms the anterior kinetosome is not ciliated. The kinetosomal dimensions measured were: length, 542 nm (508-564 nm; n=13); diameter, 192 nm (190-207 nm; n=7); and terminal plate-axosomal plate distance, 67 nm (56-71 nm; n=13). A cartwheel with a length of 128 nm (113-141 nm; n=13) was observed. Distally, four fibrillar associates accompany a kinetosomal pair. Proceeding proximally, the oblique medial desmose extends from a distal level on the anterior kinetosome to a more proximal level on the posterior kinetosome. It is from this desmose that the posterior transverse ribbon originates. The medial desmose is associated with anterior kinetosomal triplets (AKsTr) 1, 2, and 9, and posterior kinetosomal triplets (PKsTr) 3, 4, and 5. Close to the posterior kinetosome it appears denser. This electron-dense column extends proximally to form the enlarged base of desmose D1 and D2. D1 extends between AKsTr 9 and PKsTr 5 and 6, and D2 extends between AKsTr 2 and 3 and PKsTr 3 and 4. The dense material of the desmoses extends around the bases of the kinetosomes to provide anchorage for the anterior transverse ribbon, postciliary ribbon, and the kinetodesmal fibril. The same electron-dense material extends distally as two supports between the anterior kinetosome and its transverse ribbon. The kinetodesmal fibril arises externally to PKsTr 5 and 6. Its major periodicity is 28 nm (25-29 nm; n=12). It extends distally and laterally, first becoming slightly crescentic in cross section and then quite flattened and fan-shaped in the cortical ridge. Thus, in longitudinal section, this periodically striated fibril appears to taper. The other posterior kinetosomal fibrillar associates include a postciliary ribbon of 5 (3-10) microtubules (n=39) at PKsTr 9 and a transverse ribbon of 5 or 6 (3-11) microtubules (n=37) at PKsTr 3 and 4. There are usually two fibrillar associates of the anterior kinetosomes. The anterior kinetosomal transverse ribbon is composed of 10 (6-12) microtubules (n=109) arising near AKSTr 3, 4, and 5. A single postciliary microtubule is associated with triplet 9. In addition, kinetids of the right anterior and ventral regions also bear a nematodesma at the base of the anterior kinetosome. These nematodesmata originate from the dense material at the kinetosome base near AKsTr 2 and 3 and extend posteriad beneath the cortex towards the right oral polykinetid. The other kinetosomal fibrillar associates extend distally from the kinetid to become closely associated with each other and the epiplasm within the cortical ridges. The anterior transverse ribbons extend anteriad and sinistrad beneath the posterior transverse ribbons of more anterior kinetids. The posterior transverse ribbons extend several kinetids posteriad the lie laterally to each other in the ridge. Postciliary ribbons extend only a short distance posteriad and dextrad beneath the posterior transverse ribbons. Parasomal sacs, about 134 nm (92-169 nm; n=7) in diameter, are placed anteriad and posteriad of the posterior transverse ribbons and anteriad of the kinetodesmal fibril. The diameter of microtubules in all these organelles averaged 23.1 nm (20.9-24.4 nm; n=14). (ref. ID; 5918) - Oral cavity. The oral cavity is a cone-shaped depression lined by unit membrane and underlying alveoli. The latter are absent in the cytopharynx and between polykinetidal cilia. The cytostome originates in the dorsal region as a portion of the oral-cavity wall, bounded only by unit membrane. Proceeding entad, more surface area of the oral cavity is subtended by unit membrane, less by alveoli and epiplasm. This funnel-shaped region is subtended by microtubular ribbons which rotate away from the cytopharynx to form a long ribbon of microtubules, the postoral fibre. Dictyosomes are often observed in the cytopharyngeal region. Cytopharyngeal microtubules probably originate from polykinetidal kinetosomes. The ultrastructural fibrillar associates of both oral polykinetids are similar. The polykinetids are composed of rows of kinetosomes, linked by proximal longitudinal connectives. Oblique connectives link polykinetidal rows transversely. The kinetosomal dimensions were length, 574 nm (494-643 nm; n=9); diameter, 252 nm (214-268 nm; n=5); terminal plate-axosomal plate distance, 60 nm (42-85 nm; n=8); and cartwheel length, 140 nm (113-169 nm; n=3). Parasomal sacs, about 161 nm (107-214 nm; n=6) in diameter, are dispersed between rows. The terminal kinetosomes of a row bear postciliary ribbons of both convergent and divergent types. (Lynn 1976; Williams and Frankel 1973). A convergent ribbon of 7 (5-8) microtubules (n=13) is found on those kinetosomes, ontogenetically anterior, that is, closest to the cytopharynx. Phagoplasmic vesicles accompany these microtubules. Divergent ribbons of 11 or 12 microtubules (n=4) are observed on the posterior kinetosomes of a row, although some external, lateral kinetosomes do possess them. There are accompanied by dense material. Nematodesmata originate from the bases of polykinetidal kinetosomes. Overlapping each other, the extend beneath the dorsal wall of the oral cavity towards those of the other polykinetid, to form a nematodesmal bundle, possibly by interposition. (ref. ID; 5918)
- Expulsion vesicle, pore and cytoproct. The expulsion vesicle pore cannal is lined by a unit membrane underlain by a thin epiplasm. The latter is subtended by pore microtubules which number about 20 (16-21) microtubules (n=9). Radial microtubules, which extend along the expulsion vesicle surface originate in the vicinity of the pore microtubules. Occasionally an old food vacuole is observed adjacent to the anterior surface of the expulsion vesicle. Membranous components, possibly derived from these old food vacuoles, are observed adjacent to or already within the expulsion vesicle. These expulsion vesicle pore dimensions were: diameter, 1227 nm (644-1715 nm; n=13); and depth, 886 nm (466-1355 nm; n=13). A cytoproct has not been observed. (ref. ID; 5918)
- Nuclei. A highly ramified Feulgen-negative 'body' occupies the central portion of the spherical macronucleus. It is composed of densely packed nucleoli which extend to approach the macronuclear membrane. Chromatin is distributed throughout the remainder of the macronuclear volume. A single ovoid micronucleus lies adjacent to the marconucleus, often within a slight depression. Condensed chromatin is observed within it. Both macro- and micro- nuclear membranes are perforated by nuclear pores. (ref. ID; 5918)
- Cytoplasmic organelles. Food vacuoles are usually filled with bacteria. Vacuoles with freshly ingested and partially digested bacteria occasionally appear to fuse. A digestive sequence, as previously described (Rudzinska et al. 1966), can be inferred. 'Crystalline' inclusions are occasionally observed throughout the endoplasm. They are similar in appearance to those described for C. steinii (Lynn 1976). Mitochondria, whose tubules are 49 nm (39-56 nm; n=10) in diameter, granular endoplasmic reticulum, dictyosomes, and liposomes are similar to those described in the other ciliates (Pitelka 1963, 1969). (ref. ID; 5918)
- Membrane systems. A unit cell membrane covers the entire organism. It is subtended by alveoli on all surfaces with the exception of ciliary shafts and pits, expulsion vesicle pores, cytopharynx, parasomal sacs, and between the bases of oral polykinetidal cilia. The alveoli are unit-membrane-bound sacs which lie above the epiplasm. Alveolar junctions, formed by the apposition of adjacent alveolar boundaries, are filled with electron-dense material resembling that of the epiplasmic matrix. After fixations using phosphate buffer, the alveoli are indistinct. The epiplasm is a single homogeneously electron-dense layer about 26 nm in thickness, which underlies the plasmalemma or alveoli of all cortical structures with the exception of the cytostome, parasomal sacs, and the bottom of the expulsion vesicle pore. As in other Colpoda species (Lynn 1974), the epiplasm is continuous with the terminal plate of somatic kinetosomes. The cortex is ridged between somatic kineties. Anterior transverse, posterior transverse, and postciliary microtubular ribbons course within a ridge. These microtubules are closely associated with the epiplasm, perhaps by filamentous material. (ref. ID; 5918)
Measurements
35-90 um long. (ref. ID; 1618)32-40x22 um. (ref. ID; 3342)
Colpoda spiralis Novotny, Lynn & Evans, 1977 (ref. ID; 4030 original paper) reported author and year? (ref. ID; 7205)
Diagnosis
Reniform ciliate ~60-120 um long, with width ~2/3 the length; 39-55 somatic kineties; prominent somatic groove, with 9-13 kineties, spirals posteriorly and counterclockwise from oral overture; in larger individuals, prominent preoral protuberance overhangs anterior portion of oral overture; oral cavity situated in anterior 1/3 of body; vestibular kineties as extensions of 4-6 somatic kineties course along right wall of oral cavity, parallel to its right border; interior to vestibular kineties, right oral polykinetid extends along right and ventral walls; left oral polykinetid composed of 23-36 kinetosomal rows, extends along dorsal oral cavity wall; cytostome in interior dorsal part of oral cavity; macronucleus 10-30 um long, moderately ovoid, contains ramifying nucleolus; 2-14 micronuclei, 2 um in diameter; contains ramifying nucleolus; 2-14 micronuclei, 2 um in diameter; contractile vacuole pore single and posterior; division within cysts yields 2-4 tomites; resting cysts present. (ref. ID; 4030)Descriptions
- [Body shape]: When viewed from it several side, the species is reniform, with an indentation on the left lateral border. When viewed from the dorsal side, it appears less reniform, with the anterior part of the body larger than the posterior part. The somatic groove which extends over the left edge separated an anterior and a posterior portion. The body of this species is the most spiraled or twisted of the genus thus far examined in silver-stained preparations. Large specimens of Colpoda spiralis have a prominent protuberance with overhangs the anterior part of the oral cavity. Some strains averaged 120x43 um, the smaller individuals were ~40x25 um and the largest ~150x100 um. (ref. ID; 4030)
- [Kinetosome and Argyrome]: The kinetosome and argyrome of Colpoda spiralis are typically colpodid. The argyrome is a characteristic rectangular meshwork. Where the kineties converge on the posterior pole, however, this rectangular pattern becomes quite disorganized. The kineties spiral from the posterior pole in a clockwise direction (toward the organism's right), and are nearly perpendicular to the body axis on the dorsal surface. The kinetids are small and closely packed mid-dorsally in the region of the somatic groove. Anterior to the groove however, the kinetids of the kineties which traverse the groove are of a size and spacing similar to those posterior to those posterior to the groove. The kineties entering the groove at the left lateral edge continue ventrad, terminating on the left border of the oral overture. The incurrent canal kineties which course along the anterior longitudinal portion of the somatic groove, possess small anterior kinetids; the termini of these kineties do not curve posteriad. The excurrent canal kineties are separated from the incurrent canal kineties by a moderately large ridge; they are also differentiated by their larger terminal kinetids which curve slightly posterior to the previous direction of the excurrent canal kineties. Unlike those in other Colpodids, the kineties to the right of the anterior convergence region arc on the anterior right surface where they terminate, directed somewhat posteriorly. In larger individuals, the posterior extend of these kineties is correlated with the size of the preoral protuberance. (ref. ID; 4030)
- [Oral cavity]: The oral cavity is located to the right of the ventral midline and the anterior convergence region. The right border follows a strong concave curve. Extending from the right posterior broader, somatic kineties enter the oral cavity to course as vestibular kineties over its right ventral and dorsal walls. Oriented at 30-40 degrees to the long axis of the body, the left and right polykinetids are parallel in their anterior extremities. Posteriorly, the left polykinetid remains on the left dorsal wall of the oral cavity. The right polykinetid turns to course along the ventral right wall, just after is disappearance beneath the right border of the oral overture. The cytostome is located at the far right dorsal extremity of the oral cavity. Silver lines connect the borders of the oral polykinetids to each other and to the vestibular kineties. (ref. ID; 4030)
- [Contractile vacuole pore and cytoproct]: The contractile vacuole pore is located in the posterior part of the body, often slightly on the ventral surface. Silver lines connect its borders to the somatic argyrome. A cytoproct has been observed in living specimens, never in the silver-stained preparations. (ref. ID; 4030)
[Nuclei]: The moderately ovoid macronucleus may contain a highly ramified Feulgen-negative nucleolus. The remainder of the macronucleus is weakly Feulgen-positive. The length of the macronucleus ranges from 10-30 um. There are several slightly ovoid micronuclei adjacent to the macronucleus. The micronuclei, each ~2 um long, number from 2-14. (ref. ID; 4030) - [Morphogenesis after division]: The morphogenesis after division is this species is similar to that described for other species of Colpoda. Once organism have reached ~80 um in length, they form a very thin division cyst wall within which they divide. The oral apparatus dedifferentiates as the somatic kineties become bipolar. Cytokinesis ensues, resulting in 2 or 4 tomites. Stomatogenesis follows cytokinesis. It involves kinetosomal proliferation from at least 14 somatic kineties. The entire process takes 3-6 hr after which small reniform tomites excyst. (ref. ID; 4030)
[Resting cysts]: Individuals of small and medium size also form resting cysts. Larger individuals must apparently divide prior to forming the cyst. The resting or protective cyst wall is composed of at lest 2 tough, thick walls. The cyst is spherical to ovoid in shape, with an outside diameter of ~50 um from wall to wall; the encysted ciliate is spherical in shape with a diameter of ~30 um. (ref. ID; 4030)
Comments
Colpoda spiralis can be distinguished from all other congeneric species of similar shape and size. The prominent preoral protuberance present in larger specimens further distinguished members of this species. Moreover, Colpoda spiralis apparently prefers the treehole habitat. (ref. ID; 4030)Type locality
Tree hole in Quercus emoryi Torrey (Black Oak) designated N-37-66, at 31 degrees 50'N Latitude and 109 degrees 30'W Longitude, near Park Headquarters, Chiricahua National Forest, Arizona, U.S.A. (ref. ID; 4030)Holotype slide
The "type-specimens", USNM Slide no. 24489, has been deposited in the Ciliate type-collection at the Smithsonian Institution, Washington, D.C., U.S.A. (ref. ID; 4030)Colpoda steini Maupus, 1883 (ref. ID; 1335, 1622, 1629, 1796, 2245, 2342, 3116, 3593, 7533) reported year? (ref. ID; 1219, 1618) reported author and year? (ref. ID; 65, 3370, 7564), steinii Maupas, 1883 (ref. ID; 4609, 4613) reported year? (ref. ID; 3342, 3698, 3744, 4842) reported author and year? (ref. ID; 191, 7205)
Synonym
Tillina saprophila Stokes, 1884 (ref. ID; 1622, 4609, 4613)Descriptions
Body more or less kidney-shaped; mouth near lateral margin, cytostome at the bottom of a ciliated vestibulum which bears a "beard" of cilia; the margin anterior to the mouth is indented by about 5 row of cilia forming the so-called "keel"; ciliation uniform except for 2 longer caudal cilia. (ref. ID; 1219)Cytostome about two-fifths from the anterior ends, and with a bundle of long membranellae; five to six preoral ridges; paired and single cilia; one pair of long caudal cilia; twelve meridians in fresh water. The organism can live in various organs of the land slug, Agriolimax agrestis. (ref. ID; 1618)
Fine structure of the macronucleus of the active and encysted (dividing) forms. (ref. ID; 7564)
Measurements
Length 15-60 um. (ref. ID; 1219)15-42 um long. (ref. ID; 1618)
20-30x10-13 um. (ref. ID; 3342)