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The Stalk of a Peritrich Ciliate Vorticella Is Usually 0.2-0.5Mm Long and Can

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The Stalk of a Peritrich Ciliate Vorticella Is Usually 0.2-0.5Mm Long and Can J. Biochem. 83, 795-798 (1978) Improved Preparation and Cooperative Calcium Contraction of Glycerinated Vorticella1 Hiroshi ASAI, Tsutomu OCHIAI, Keiji FUKUI , Masato WATANABE, and Fumio KANO2 Department of Physics, Waseda University, Nishi-okubo, Shinjuku-ku, Tokyo 160 Received for publication, September 9, 1977 An improved method for the preparation of glycerinated Vorticella convallaria was investigated. The pretreatment of living vorticellas with a medium containing 0.1 % saponins and subsequent treatment with an extraction medium containing 35 % glycerol at about 0•Ž was satisfactory. The equilibrium average length of contractile stalks of glycerinated vorticellas was measured at various free calcium concentrations in the reaction medium. It was found that the contractile element in the spasmoneme of the stalk is contracted by a cooperative interaction involving at least two calcium ions. The stalk of a peritrich ciliate Vorticella is usually times encountered a similar problem, which is 0.2-0.5mM long and can coil into a left-handed probably mainly due to disconnection of the helix. The coiling is produced by the contraction contracted spasmoneme from the sheath. Thus, of an intracytoplasmic thread called the spasmo we sought to develop an improved method for the neme, which is coiled spirally within a cylindrical preparation of glycerinated Vorticella. sheath of the stalk. Hoffmann-Berling (1) demon It was found after several trials that the strated that the stalk of glycerinated Vorticella pretreatment of living vorticellas with 0.1 coiled in the presence of calcium ions and uncoiled saponins followed by gradual glycerination is when calcium was subsequently removed by the satisfactory. This report describes this improved addition of a calcium chelater, e.g., EGTA. preparation method and the cooperative properties However, Townes and Brown (2) reported of calcium-contractile Vorticella. that the reversible contraction of glycerinated preparations by calcium ions becomes weak, MATERIALS AND METHODS especially at pH's far from pH 6.8. We some The vorticella used was a clone isolated from a 1 This work was aided in part by the Saneyoshi Science population taken from a local pond and identified Foundation. as V. convallaria. Stock cultures of the clone 2 Present address: Tokyo Metropolitan Laboratory of were maintained on cover glass fragments in hay Public Health. infusion medium previously inoculated with Aero Abbreviations: EGTA, ethyleneglycol-bis(P-amino-ethyl bactor aerogenes as prey for the vorticellas. The ether) N,N•Œ-tetraacetic acid; EDTA, ethylenediamine cover glass fragments with vorticellas attached to tetraacetic acid. Vol. 83, No. 3, 1978 795 796 H. ASAI, T. OCHIAI, K. FUKUI, M. WATANABE, and F. KANO both surfaces were transferred from the culture medium to a medium containing 0.1 % saponins, RESULTS AND DISCUSSION 4mM EDTA, 100mM KCl, and 20mM histidine HCl buffer (pH 6.8) at room temperature. The When living vorticellas cultured in hay infusion medium was cooled gradually using an ice-cold medium were immersed directly in 50% glycerol, water bath and kept for 45 min at this temperature. the vorticellas were very often coiled tightly so The gradual cooling of the medium in which the that the spasmoneme became detached from the cover glass fragments with vorticellas were placed sheath and lost its reversible contractivility. The was essential for obtaining good preparations. bell-shaped zooid was also easily detached from The cover glass fragments were then immersed in the sheath. Thus, it was important to glycerinate a new extraction medium containing 35% glycerol, the living vorticellas by means of a gradual increase 4mM EDTA, 100mM KCl, and 20mM histidine of glycerol concentration. It was also important HCl (pH 6.8) at about 0•Ž. The extraction was to change the temperature of the extraction medium continued for 60 min at this temperature. For or reaction mixture gradually. After several prolonged storage (up to 6 months) of glycerinated trials, it was found, however, that the glycerinated preparations of vorticellas attached to the cover models thus prepared still sometimes lost the glass fragments, the extraction medium was ability to exhibit reversible contraction-extension replaced by a new extraction medium containing cycles if the pretreatment with 0.1 % saponins was 50 % glycerol, 4mM EDTA, 100mM KCl, and 20 omitted, and if EGTA rather than EDTA was mm histidine-HCl buffer (pH 6.8) and stored at used as a Ca-buffer in reaction mixtures. Of -15 -20•Ž. For immediate use of glycerinated course, the use of EGTA is essential to adjust preparations, 35 % glycerol was removed by rinsing free calcium concentration in a reaction mixture with a solution containing 4mM EDTA, 100mM containing free magnesium ions. After pretreat KCl, and 20mM histidine-HCl (pH 6.8) at about ment of living vorticellas with 0.1 % saponins, the 0•Ž for 10 min. Then, the rinsing medium was subsequent treatment with 35 % glycerol for 60 min warmed gradually to room temperature and a was sufficient to obtain excellent models, as de cover glass fragment with attached vorticellas was scribed in the previous section. It was found, placed in a well on a slide glass containing a during the course of various experiments, that for suitable reaction mixture in the presence of calcium immediate use the treatment of living vorticellas buffer. For the use of stored preparations of with 0.1 % saponins alone for 2 h was sufficient. glycerinated Vorticella in 50% glycerol, the glycerol Actually, Hofmann-Berling (1) stated in his paper was removed by rinsing with a solution containing that 0.1 % saponins can be used to pretreat living 4mM EDTA, 100mM KCl, and 20mM histidine Vorticella gracilis without mentioning the effects HCl (pH 6.8) at about 0•Ž for 10 min. Then, of saponin pretreatment on the vorticellas. It is the rinsing medium was warmed gradually to room generally considered that one of the effects of sapo temperature. The subsequent procedures for ex nin treatment of a biological membrane is to en periments were similar to those already described. hance the permeability of the membrane to various All reaction mixtures for measuring the ions and also to some macromolecules (5). How stalk lengths of vorticellas contained 4mM EGTA ever, subsequent investigators of glycerinated vorti as the calcium buffer, 100mM KCl and 20mM cellas, i.e. Townes and Brown (2), Amos (4), and histidine-HCl (pH 6.8). A value of 5 x 105 m-1 others, have not followed Hofmann-Berling's pre was used as the stability constant of EGTA and treatment method. Thus, our method of preparing calcium ions to estimate pCa at pH 6.8 (3). The glycerinated vorticellas is essentially a redevelop stalk lengths of vorticellas attached to a cover ment of his method. glass fragment on the slide glass were measured The contraction and extension cycles of two by means of an ocular micrometer at a magnifica glycerinated stalks thus prepared are shown in tion 40 x or 100 x after each change of concentra Fig. 1. The cycles were induced more than 135 tion of free calcium. times by immersing the stalks alternately in reaction solutions containing 1ƒÊM free calcium (estimated) and 4mM EGTA. The fractional stalk length J. Biochem. PREPARATION AND CONTRACTION OF GLYCERINATED Vorticella 797 (L) is taken as 1.0 for the fully extended state and chemical one and does not involve ATP or other as 0.0 for the completely contracted state. As glycerol-extractable chemicals. shown by the broken line in Fig. 1, the ability to When the free calcium concentration in the contract in 1ƒÊM free calcium gradually declined reaction mixture was changed from 10 pat to with repeated cycles of contraction and extension. 2 [LM, the stalk extended partly, as shown im As shown by the cross in this figure, the addition Fig. 2, and maintained its extended state for more of 1mM ATP did not affect the contractibility of than 1 h. The time required to extend after the stalks. The results confirm that the contrac changing the medium was less than I min. The tion of the vorticella stalk is a purely mechano required time to contract after changing to a medium containing a higher concentration of free calcium ions was similar. It is thus con sidered that during contraction and extension cycles the equilibrium stalk length is reached within 1 min. The average fractional stalk length of many vorticellas is plotted against the free calcium concentration in reaction mixtures in Fig. 3. At free calcium concentrations lower than 0.1ƒÊM, the glycerinated stalks are fully extended. At free calcium concentrations higher than 10ƒÊM, the stalks are invariably contracted. The frac tional stalk length (L) can be expressed as a func tion of free calcium concentration by a Hill's Fig. 1. Plot of many reversible contraction-extension equation, which is frequently applied to ligand cycles of glycerinated Vorticella convallaria. The reac binding with an allosteric protein, tion mixtures for the contracted states of one stalk (ƒ•) and another stalk (ƒ¢) contained 100mM KCl, 20mM histidine-HCI (pH 6.8), 4 mat EGTA, and 1ƒÊM (esti mated) free calcium ions. (•~) indicates the presence of 1mM ATP in the above reaction mixture. The reaction mixture for the fully extended state of the stalks (•œ) contained 100mM KCI, 20mM histidine-HCI (pH 6.8), and 4mM EGTA. Fig. 2. Time courses of stalk length in various free calcium concentrations. Vol. 83, No. 3, 1978 798 H. ASAI, T. OCHIAI, K. FUKUI, M. WATANABE, and F. KANO Fig. 3. Relationship between the normalized overall stalk length and pCa. A logarithmic plot of (1-L)/L against pCa is shown in the insert for estimation of the Hill's parameter n and affinity Km. where [Ca] is the concentration of free calcium in the coiled-stalk length after full coiling was more ions and pCa=-log10[Ca].
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