Blepharisma

Blepharisma Perty, 1849 (ref. ID; 2014) or 1852 (ref. ID; 3690)

Class Polyhymenophora: Subclass Spirotricha: Order Heterotrichida: Suborder Heterotrichina (ref. ID; 2014)

[ref. ID; 480]
The classification of Blepharisma is most usually based upon nuclear characteristics (Hirshfield et al. 1965; Suzuki 1954) i.e., macronuclear shape and behavior during conjugation and micronuclear number and size. Other characteristics, such as body size, peristomal length, number of somatic ciliary rows, membranelle number of the AZM are also useful morphological details. (ref. ID; 480)

[ref. ID; 2014]
Body elongate and lenticular, size varies from medium (50 um long) to very large (1 mm long), anterior bluntly pointed with slightly curved lip. Terminal pole rounded. Body is non-contractile but variable in size and shape even within clonal cultures, slightly flattened when underfed. Uniform ciliation but the number of ciliary rows depends on size of animal not on species. Pigmentation common, usually pale pink to bright red with pigmentation granules in longitudinal bands alternating with ciliary meridians. Oral apparatus along left edge winding onto ventral surface and consisting of AZM on the left of the peristomial border, a conspicuous undulating membrane on the right and a concave groove between the two. Single large posterior contractile vacuole without collecting canals. Macronucleus may be rounded, elongate moniliform or be composed of several spherical macronuclei. Genus most easily mistaken for Spirostomum which is rarely colored (never pink or red), is contractile and has a collecting canal serving the contractile vacuole.
Quote; Colin R. Curds, Michael A. Gates and David McL. Roberts "British and other freshwater ciliated protozoa Part II Ciliophora: Oligohymenophora and Polyhymenophora" Cambridge University Press, 1983 (ref. ID; 2014)

Blepharisma americanum (Suzuki) (ref. ID; 646, 1308) reported author and year? (ref. ID; 4025, 4429)
Blepharisma arctica (ref. ID; 3695)
Blepharisma bimicronucleata Villeneuve-Brachon (ref. ID; 4842) reported author and year? (ref. ID; 191)
Blepharisma botezali Lepsi, 1926 (ref. ID; 1621)
Syn; Blepharisma lateritium var. botezali Lepsi (ref. ID; 1621)
Blepharisma bothrostoma Mermod, 1914
See; Metopus undulans (ref. ID; 1621)
Blepharisma candidium Yagiu & Shigenaka, 1956
See; Anigsteinia candidium
Blepharisma clarissimum Anigstein, 1912
See; Anigsteinia clarissimum (ref. ID; 1621)
Blepharisma clarissimum f. arenicola Kahl, 1932 (ref. ID; 3119), var. arenicola Kahl, 1932 (ref. ID; 3690)
Blepharisma clarissimum var. longissimum (ref. ID; 1621)
Syn; Blepharisma clarissimum Kahl, 1928 (ref. ID; 1621)
Blepharisma coeruleum Gajevskaja, 1927 (ref. ID; 1621, 1629, 4610)
Blepharisma dileptus Kahl, 1928 (ref. ID; 1621)
Blepharisma elongatum Stokes, 1884 (ref. ID; 1621, 3593, 3694) reported year? (ref. ID; 3698)
Blepharisma galianoi (ref. ID; 4429)
Blepharisma grayi Hartwig & Parker, 1977 (ref. ID; 2316)
Blepharisma hyalinum Perty, 1849 (ref. ID; 4861) or 1852 (ref. ID; 1308, 1335, 1621, 3593, 4057) reported year? (ref. ID; 3698)
Syn; Blepharisma lateritium var. minima Roux (ref. ID; 1621)
Blepharisma ichthyoides V. Gelei, 1933 (ref. ID; 644, 1620)
Blepharisma intermedium (ref. ID; 4057)
Blepharisma japonicum Suzuki (ref. ID; 4017, 7544) or (Suzuki, 1954) Giese, 1973 (ref. ID; 1618) reported author and year? (ref. ID; 191, 4025, 4900, 4901)
Syn; Blepharisma undulans japonicus (ref. ID; 1618)
Blepharisma japonicum var. intermedium (ref. ID; 4025, 7544) Blepharisma lateritia (ref. ID; 191)
Blepharisma lateritium (Ehrenberg, 1831) (ref. ID; 1335, 1620, 1621, 1629, 2245) reported year? (ref. ID; 1219, 1618, 3698), (Ehrenberg, 1831) Kahl, 1932 (ref. ID; 644) or (Ehrenberg, 1831) Stein, 1859 (ref. ID; 4610) reported author and year? (ref. ID; 4429)
Syn; Blepharisma lateritium var. navicula Lepsi, 1926 (ref. ID; 1621); Bursaria lateritia Ehrenberg, 1831 (ref. ID; 4610)
Blepharisma lateritium var. bucoviniense Lepsi, 1926 (ref. ID; 1621)
Blepharisma lentis (ref. ID; 4610)
Blepharisma minima Lepsi, 1926
See; Protocrucia adhaerens (ref. ID; 1621), Protocruzia contrax (Mansfeld, 1923) (ref. ID; 4905)
Blepharisma multinucleata Dragesco, 1960 (ref. ID; 3690 original paper)
Blepharisma musculus Penard, 1922 (ref. ID; 1308, 1621)
Syn; Apgaria undulans Stokes, 1884 (ref. ID; 1621)
Blepharisma musculus var. seshachari Bhandary, 1959 (ref. ID; 508)
Blepharisma ovatum Kahl, 1927 (ref. ID; 1621)
Blepharisma ovatum Penard, 1922 (ref. ID; 1621)
Blepharisma persicinum Perty, 1852 (ref. ID; 480, 1621) reported year? (ref. ID; 1618)
Blepharisma salinarum Florentin, 1899 (ref. ID; 1621) reported year? (ref. ID; 3771)
Blepharisma sphagni Lepsi, 1926 (ref. ID; 1308, 1621, 3593)
Syn; Blepharisma lateritium var. sphagni Lepsi (ref. ID; 1621)
Blepharisma steini Kahl, 1932 (ref. ID; 3593) reported year? (ref. ID; 1308, 1618, 1620, 1621, 3690) reported author and year? (ref. ID; 4057, 4429)
Syn; Blepharisma lateritium Claparede & Lachmann (ref. ID; 1621)
Blepharisma stoltei Isquith (ref. ID; 4111)
Blepharisma tardum Kahl, 1928 (ref. ID; 1621)
Blepharisma tenuis Kahl, 1926
See; Pseudoblepharisma tenue (ref. ID; 4610)
Blepharisma undulans Stein, 1868 (ref. ID; 1308, 1621) or 1867 (ref. ID; 2288) or reported year? (ref. ID; 1618, 4429) reported author and year? (ref. ID; 191)
Blepharisma undulans undulans Suzuki, 1954 (ref. ID; 1618)
Blepharisma undulans americanus Suzuki, 1954 (ref. ID; 1618)
Blepharisma undulans japonicus Suzuki, 1954 (ref. ID; 1618)
Blepharisma vestitum (ref. ID; 1621)
Blepharisma wardsi (ref. ID; 2346)

Blepharisma elongatum (Stokes, 1884) (ref. ID; 1621, 3593, 3694) reported year? (ref. ID; 3698)

Descriptions

B. elongatum has a compressed shape which is asymmetrical in longitudinal section; the left side is flat or slightly curved, whereas the right side is more or less convex, dependent on the number of food vacuoles present. The swimming behaviour of the ciliate was in the form of a spiral, curving to the left. The "tail" is ciliated and the contractile vacuole is positioned anteriorly to this anatomical details. The peristome extends about half the body length and it contains an adoral zone of about 40 membranelles (AZM) and a conspicuous undulating membrane (UM) extending about half the length of the peristome. The macronucleus is compact and oblong and the micronuclei close to, or in indentations of, the macronucleus. This species colorless. The somatic region of B. elongatum revealed structures typical of Blepharisma. Hence the somatic ciliature was composed of dikinetids of which only one kinetosome was ciliated and of kinetodesmal fibers, composed of postciliary fibrils from the dikinetids, passing along the kineties. Between the kineties were pellicular ridges below which a regular pattern of pellicular vacuoles could be seen. The pellicle seemed to consist of two closely associated membranes with an electron dense layer on the outside. However, a characteristic feature of the somatic region was the almost total absence of cortical granules, only occasionally were such granules seen, but extrusion of their contents was observed. The oral region revealed a typical undulating membrane of closely packed dikinetids of which only one kinetosome was ciliated. A large portion of the peristome was occupied by cytopharyngeal ribbons. The adoral zone of membranelles (AZM) was found to be composed of membranes containing 3 or 2 rows of kinetosomes. The narrower membranelles were found closest to the cytostome, a feature which could also be revealed in protargol preparations. The nematodesmata, arising from two kinetosomal rows of the membranelles, passed far into the cytoplasma and showed a typical hexagonal packing of microtubules. Unlike the somatic region, the oral region contained several cortical granules; some granules had contents of fluffy material resembling the few granules seen in the somatic region, whereas the contents of other granules revealed arrays of electron dense material. Although the latter type of granules was often seen as oblique structures, they never appeared as the typical peristomal rods found in other Blepharism species. The macronucleus contained condensed threads of chromatin material and interspersed nuclei. The highest number of micronuclei seen in sectioned ciliates was two, they were positioned in macronuclear indentations of which part was occupied by cytoplasm containing rough endoplasmic reticulum. The nuclear envelopes of the macronucleus and micronuclei were typical double membranes with nuclear pores. The general cytoplasm of the ciliates was highly vacuolated, but also food vacuoles were observed. The mitochondria were of the tubular type. Some of the ciliated also contained several paraglycogen bodies. (ref. ID; 3694)
[Cyst]: The cysts of B. elongatum were typical of the Blepharisma genus. The cyst wall was composed of two layers, the outer shell, or ectocyst, and an inner membrane, or endocyst; the emergence pore was well developed. The compact macronucleus was recognizable in the native cysts, but micronuclei could not be detected. (ref. ID; 3694)

Measurements

215 um long and 65 um wide and its macronucleus measured 50 by 20 um. (ref. ID; 3694)

Blepharisma grayi Hartwig & Parker, 1977 (ref. ID; 2316)

Descriptions

This specimen is displaying four membranelles of long cilia on the right margin of the anteriormost quarter of the peristome, together with an undulating membrane. Three large vacuoles and several smaller ones visible near the dorsal surface. (ref. ID; 2316)

Measurements

Length 170 um. (ref. ID; 2316)

Blepharisma hyalinum Perty, 1849 (ref. ID; 4861) or 1852 (ref. ID; 1308, 1335, 1621, 3593, 4057) reported year? (ref. ID; 3698)

Synonym

Blepharisma lateritium var. minima Roux (ref. ID; 1621)

Descriptions

Blepharisma hyalinum was colorless when originally isolated from nature in 1975 and whenever it has been recollected. The organisms remains colorless growth in the dark in laboratory culture. B. hyalinum isolated from nature appears slender and flexible, whereas organisms from cultures are typical pyriform in shape. The latter organism have an average length of 100+/-20 um (n=50), with range 80-160 um, and an average width of 36 um, range 24-48 um. The somatic ciliature consists of ca. 18 kineties situated in cortical depressions. The peristome extends about half the body length with an AZM of ca. 30 membranelles and a conspicuous undulating membrane extending about half the length of the peristome. The compact, oblong macronucleus is ca. 25x12 um. Some 4-7 micronuclei, ca. 2 um in diameter, are located near the macronucleus. The large contractile vacuole, typical of Blepharisma, is situated posteriorly. Even though B. hyalinum is colorless, it contains cortical granules, visible under phase contrast optics; the granules are arranged in bands between the kineties and each band has a width of 4-5 granules. On addition of alcian blue, the granules extrude their contents, forming an intensely stained "shell" around the organism like that described for Tetrahymena (Nilsson, 1972). (ref. ID; 4057)

Blepharisma ichthyoides V. Gelei, 1933 (ref. ID; 644, 1620)

Comments

B. ichthyoides 150 um in length, 30-45 um in width, the colour coppery red and the posterior portion of the body cylindrical with the terminal end swollen near the contractile vacuole. However, Gelei was note quite sure of its validity and considered that B. lateritium, B. steini and B. ichthyoides might be variations of one species. The present investigation seems to indicate, that B. ichthyoides in fact is the pre-encystment stage of B. lateritium. (ref. ID; 644)

Blepharisma lateritium (Ehrenberg, 1831) (ref. ID; 1335, 1620, 1621, 1629, 2245) reported year? (ref. ID; 1219, 1618, 3698) or (Ehrenberg, 1831) Kahl, 1932 (ref. ID; 644), (Ehrenberg, 1831) Stein, 1859 (ref. ID; 4610) reported author and year? (ref. ID; 4429)

Synonym

Blepharisma lateritium var. navicula Lepsi, 1926 (ref. ID; 1621); Bursaria lateritia Ehrenberg, 1831 (ref. ID; 4610)

Descriptions

The body of B. lateritium is usually pyri-to dropdormed with the anterior end pointed and a little flattened, while the posterior end is expanded and almost circular in cross section. The buccal apparatus is commonly 2/3 to 3/4 the length of the whole organs. It is furnished on the lower part of the right margin with an undulating membrane with a length of about 40 um and a width of about 10 um. The left side of the oral groove is furnished with a well developed adoral zone of 80-90 membranelles which together with the undulating membrane leads down to the cytostome, where the food vacuole are formed and transported into the cytoplasm assisted by an arrangement of conspicuous pharyngeal fibrils. The body ciliature is arranged in about 40 rows dispersed in 20 on each side. The rows of cilia on the right side run parallel to the peristomal margin, while those on the left side are forming an angle of about 60 degrees with the left margin of the peristome. By analogy with other strains of Blepharisma, the colour of B. lateritium is variable dependent on different environmental factors. Organisms collected from the nature are usually pale pink to nearly colourless, while those grown in the darkness in cultures become more deeply pigmented. The pigment granules which are considerably smaller (about 0.25 um) than those from other forms, e.g. B. persicinum, are arranged in between the kineties in rows of 14-16 granules across. The macronucleus which is spherical to body and about 25-30 um in diameter, is centered in the anterior half of the body. There are 4-8 small micronuclei with a diameter of about 2 um. There is one large contractile vacuole located terminally, where also the cytopyge is placed. Although this organelle may expand enormously during the defecation procedure, no permanent excretion pore has been observed. (ref. ID; 644)

Binary fission: The first sign of division is observed with a change of the characteristic drop form into a more "rhomboid" form of the body. At the same time a new contractile vacuole and a new AZM anlage is established posteriorly to the old oral area. During this stage phase no visible change of the macronucleus is observed. As a primary compact macronucleus is already present, there is no condensation phase prior to the division and the macronucleus just elongates until it is divided between the two daughter cells, thus the binary fission process is analogue to that of most other ciliate species. The micronuclear mitosis of most Blepharisma species is commonly correlated with the macronuclear division, but although numerous division stages have been examined, no micronuclear division stages have been observed. An explanation to this may be that the micronuclear mitosis possibly begins before any sign of division is visible and that such early stages simply have been overlooked. (ref. ID; 644)
Conjugation: Conjugation stages or at least pairs of attached cells have been observed only on organisms collected from their cool natural environment and placed at room temperature in the laboratory. Conjugation has never been observed in established laboratory cultures. Although several conjugating pairs have been examined, the conjugation process seems not to have been completed, as stained preparations of the cells never showed signs of macronuclear reorganization. On the other hand the micronuclei were active during the process and different stages of micronuclear pregamic divisions were observed. (ref. ID; 644)
Cyst: Encystment has frequently been observed in collections brought from the cold in the nature into room temperature, but is also occurs commonly in old laboratory cultures. In its vegetative stage B. lateritium is very slow in its movements and in cultures with abundant food supply the organisms very often use to keep still and just whirl in food organisms, bacteria and small algae, by the movements of its oral apparatus. Prior to encystment, however, drastic changes are taking place. The first sign of a beginning encystment is a lively swimming around as if the organisms were seeking a hiding-place, and in fact they are. In the culture vessels they very commonly accumulate in great numbers underneath the wheat kernels, where the final encystment process takes place. Together with this increase in activity striking cytological changes take place. The body assumes a lancet-form with the posterior portion thin and cylindrical and with the terminal end swollen near the contractile vacuole. The peristome ceases to ingest food and the oral organelles begin or de-differentiate. The food vacuoles disappear presumably with a loss of water and an increase in the density of the cytoplasm. A simultaneous aggregation of the pigment granules makes the organisms appear a deeper red in colour. At the same time the macronucleus seems to undergo a reorganization and finally the organisms settle down, contract to a regular circular shape, the pre-stage of the final encystment process. The nuclear pattern of the resting cyst seems to be identical with that of the vegetative organism. Although numerous experiments have been performed, excystment could so far not be induced under laboratory conditions. (ref. ID; 644)

Pyriform; macronucleus oval; a micronucleus; rose-colored; fresh water among decaying leaves. (ref. ID; 1618)

Measurements

The average length of the organism is 160+/-16 um (N=50) ranging from 110 to 190 um. The average width is 90+/-11 um with a range from 70-110 um. (ref. ID; 644)
130-200 um long. (ref. ID; 1618)

Blepharisma persicinum Perty, 1852 (ref. ID; 480, 1621) reported year? (ref. ID; 1618)

Descriptions

The species appears flattened and highly transparent the whole body though the posterior part may be more or less rounded according to the number of food vacuoles present. The somatic ciliature is dense and uniform. About 20 rows of cilia, kineties, on the right side of the organisms run anterior-posterior, parallel to the oral groove, while on the left side they meet the groove at an angle of about 45 degrees, the number of kineties on this side being about 10. The buccal apparatus is approximately 2/3 the length of the whole organism. It is provided on the lower part of the right margin with an undulating membrane (UM) which is composed of the cilia arising from the posterior third of the peristomal margin. The membrane can easily be observed while the cell is alive, but breaks up into is component cilia during fixation and staining. The entire left side of the groove is furnished with an adoral zone of about 60 membranelles (AZM), which together with the UM posteriorly leads down the funnel shaped part of the oral area to the cytostome (CY), where the food vacuoles (FV) are formed. The contractile vacuole (CV) is located posteriorly, where also the cytopyge (CYP) is placed, but no permanent pore has been observed. The vegetative macronucleus (MA) is the present strain of Blepharisma consists of 4-9, generally 7, spherical nodes arranged as a bent string of beads with diameters from about 5 to 10 um. No connective strands are revealed neither by phase contrast nor by light field microscopy of stained cells. The number of micronuclei (MI) is between 4 and 7 during interphase; each micronucleus is about 3 um in diameter, considerably larger than those reported from other strains of Blepharisma. (ref. ID; 480)

Binary fission: The sequence of macronuclear events during binary fission of Blepharisma is not always the same, and depends on the macronuclear configuration of the species. When a primary compact macronucleus is already present, it elongates until it is divided between the two daughter cells and then contracts to assume the interphase shape. In Blepharisma's with a multinodal macronucleus a condensation of the latter takes place to form a spherical macronucleus, which reelongates before the cell splits to two daughter cells. The micronuclear division is correlated with the macronuclear division and the micronuclear mitosis begins after the macronucleus contracts into its spherical from and is completed when the fully condensed macronuclear mass has nearly completed its reelongation. Conjugation has been observed in the cultures but so rarely that this process could not be studied in detail. (ref. ID; 480)
Cyst: Like many free living ciliates Blepharisma is also able to form cysts, but the frequency of encystment varies from species to species. A wide variety of conditions have been regarded as favoring encystment, for example: lack of food, the accumulation of excretory products, changes in hydrogen-ion concentration, evaporation of the medium etc. The formation cysts in B. persicinum has been observed particularly in old cultures, but details in the encystment and excystment process have not been studied. However, fixation and staining of the cysts have revealed the same configuration of the nuclear material as present in the vegetative stage of the organism. (ref. ID; 480)
Pigmentation: Perhaps the most interesting feature of Blepharisma is its possession of Blepharismin a photosensitizing pigment somewhat like Hypericin, but in view of the fact that Blepharisma can be completely bleached by exposure to light, the color of field specimens can not be used for determining classification. The pigmentation of B. persicinum is similar to that of other strains, though the pigment granules seem more irregularly scattered between the rows of cilia as seen for instance in Blepharisma japonicum. Like other strains B. persicinum also becomes more deeply pigmented when grown in laboratory cultures in the dark. (ref. ID; 480)

Elongate oval; posterior end pointed; left peristomal edge sigmoid; preoral membrane large; macronucleus in 3-7 parts; rose-colored; fresh water among decaying vegetation. (ref. ID; 1618)

Comments

It was temporarily identified as Blepharisma persicinum, which was first described in 1849 by Maximillian Perty, who in 1852 created the genus Blepharisma (Perty 1849, 1852). This first description was short and fairly incomplete. The description was illustrated by a drawing, but it was insufficient for a later identification and although B. persicinum has been redescribed by later investigators (Penard 1922; Kahl 1932; Dingfelder 1962; Jankowski 1964) it has been mentioned as a nomen nudum as late as in 1973 (Giese 1973). Wilfert (1972) in his thorough survey on the taxonomy of the genus Blepharisma claimed that previous investigators Penard (1922); Kahl (1927, 1932) and Dingfelder (1962) have mentioned 3-7 macronuclear nodes but that they have overlooked the strands connecting them. However, connective strands have not been observed during the present investigation, neither by phase contrast microscopy of living cells nor by the examination of fixed and stained material. Further Wilfert emphasizes the taxonomic position of B. persicinum as uncertain, but he is pointing out that the diagnosis of Penard must still be considered at the most valid of the earlier descriptions, a point of view that is in good agreement with the present investigation. (ref. ID; 480)

Measurements

The average length of the organism is about 125 um with a range from about 95 to 145 um. The average width is about 65 um ranging from about 50 to 70 um. (ref. ID; 480)
80-120 um long. (ref. ID; 1618)

Blepharisma steini Kahl, 1932 (ref. ID; 3593) reported year? (ref. ID; 1308, 1618, 1620, 1621, 3690) reported author and year? (ref. ID; 4057, 4429)

Synonym

Blepharisma lateritium Claparede & Lachmann (ref. ID; 1621)

Descriptions

Macronucleus ovoid; reddish to colorless; fresh water in sphagnum. (ref. ID; 1618)
Blepharisma steini becomes pigmented after growth in the dark. It has an average length of 146+/-17 um (n=50), with range 100-184 um, and an average width of 52 um, range 44-60 um. The somatic ciliature consists of ca. 25 kineties. The peristome extends about half the body length; the AZM contains ca. 50 membranelles and the undulating membrane extends about a third of the length of the peristome. The compact, spherical macronucleus has a diameter of ca. 20 um and there are 4-7 micronuclei, each ca. 2 um in diameter. The pigment granules are arranged in bands, 7-10 granules across, between the kineties. Light-bleaching of B. steini results in a considerable reduction in number of granules as observable under phase contrast optics. Furthermore, formation of the intensely stained "shell" around the organisms, the presence of alcian blue, is accompanied by loss of pigmentation. (ref. ID; 4057)

Measurements

80-200 um long. (ref. ID; 1618)

Blepharisma stoltei Isquith (ref. ID; 4111)

Descriptions

Resting cysts: Section of thick-walled cyst, inner wall enclosing protoplasm and plug. (ref. ID; 4111)

Blepharisma undulans Stein, 1868 (ref. ID; 1308, 1621) or 1867 (ref. ID; 2288) reported year? (ref. ID; 1618, 4429) reported author and year? (ref. ID; 191)

Descriptions

Macronucleus in 2 parts; undulating membrane long; cytopharynx directed posteriorly; fresh water among decaying vegetation. Suzuki (1954) distinguishes three subspecies, based on the form and behavior of the macronucleus during division; B. u. undulans, B. u. americanus and B. u. japonicus. (ref. ID; 1618)

Measurements

150-300 um long. (ref. ID; 1618)