STUDY ON CYCLOSIS OF PARAMECIUM
Akira
(Biological
Institute,
(1)
KOENUMA*
Faculty
of Libetal
Aits and:Science)
Introduction
The cyelosis.of Paramecium was discovered iri 1836,in P. bursaria by
FOCKE, and then the existence of cycloSis.in P.:cciudatum.or P. aurelid
was brought out clearly. But in spite of the old discovery, 'considerably
few observations on this phenomenon were reported, and at present there
are the reports of observations performed by WALLENGREN (1902),
NIERENSTEIN (1905), BILLS (1922),and HOSOI (1936). The interesting
fact on motive force of the cyclosis of •paramecium was reported by
HOSOI in 1937.
A quite convenient method to stop the P.rameciiim was .discovered
by BILLS in 1922. This BILLS' narcotizing method by isopropyl alcohol
Was very effective to stop Paramecium without injuring the endoplasm;
and in HOSOI's experiment on cyClosisof Paramecium, a method based
on the BILLs' narcotizing method was used.
According to the observations carried out before', cyclosis of P
caudatum is as follows. If one assumes that the side where the oral
groove is present is the "ventral" of Paramecium, the cyclosis flows
counter-clock wise when Paramecium is observed from ventral side. It
flows to anterior direction in left of the body, and to posterior direction
in right of the body. It occupies almost the whole space of endoplasm,
and the boundary between the streaming and the rest is obscure. In
side view, cyclosis flows to be 00-shape, because it flows along the
inner surface of oral groove at the left anterior part, then flows along
the inner surface of oral fold,' then streaming removes to the dorsal
portion of the body at cytopharynx, at last it removes to the ventral
portion of the body at posterior end of the body. Any reversal of the
direction of cyclosis has not been observed yet.
This is one of the papers on cyclosis of Patamecium, in which the
effects of the body-length, temperature and concentration of external
medium, and K-, Na-, Ca- and Mg- ions in the medium were descrived
and discussed.
Material
Paramecium
hay
*
culture,
Assistant
was
and
their
of Shinshu
the
caudatum
body-length
University
and
type
were
.
Method
which
between
was
obtained
16011 and
21011.
from
the
Before
A． KOENUMA
50（ 2 ）
’ No． 4
the observations they were made to adapt to the test solutions for 18
to 20 hours after being washed in 100 volumes of redisti11ed water to
one volume of culture medium． The pH of the test solutions were kept
between 7．1・ and 7．3 by addition of Na｝ICQ3． At the observation of．
cyclosis， either of the following two methods was used to stop the
Pavamecium： a narcotizing method by isopropyl alcohol vapour， which was
used by HOSOI， and a new method in which viscous medium， 1％ traga−
cantha solution， ， was’ used together with the narcotizing by isobropyl al−
cohQl vapour． By using the latter， a good result was obtained．
The velqcity， of cyclosis was obtained by measuring the time requi・；
red． fQr the granules． in cyclosis to． flow fQr 15Lt． near of oral groove of
PaPta郷eciM解．
ebseivvations and Discinssion
1） Effect of the body−length of Paptamecium on cyc］osis．
The re＄ult of the measurement of velocity Qf cyclosis of Parame−
cium which had the body−length of from 160pt to 210pt，． was indicated in
table 1 and fig． 1， The． test solution； pH 7．3 of artificial sea water was
used， and the temperature was 260C． ・The concentration of the test
solution was equivalent to O．02 M NaCl・ solution．
Table 1
The body−iength en cyclosis of Parameciasm．
Remarl〈s ：
body−length
Velocl’ty of・ cyclosis
16． 0−170コ口
2．gpt／s
170−180
2．9
・180−190
B．0
190−200
200−210
3．0
2．9
260C
Temperature
Solution
O．02 Mol artificia1 sea wate’r pH＝7．3．
AdaptatiQn
20 ho，urs．
4
鱒劔
⑲
二面
四鰍
⑤爾
刻⑳畢
㊥
闘⑱
㈱
幽
B
？
西ぢ2①﹀
1
0
160 170 180 190 200 210P
Body−inngth
Fig． 1 T．he body−length on cyclesis ef Parameci’ 浮香D
Thb velocjty of’ cyclosis and ’ 狽??@body−Jength were indj’cated on the
ordinate and the absissa of the graph respectively．
Each circle represents the velocity．
No． 4
STUE）Y ON CYCLOSIS OF PARAMECIUM （1）
（ 3 ）51
In負g．1， the velocity 6f cyc工osis ahd the bodyL工ength were ihdic畿ed
on the ordinate and on the absiSsa of the graph resp’ectiV’ely． ’Each， citele・
on this graph represents the veloclty of cyclosis．・ ln these−table and／
且9．，童tis apParent that the ve16citY of cyc工osis is not d◎ncerned Svith・the
body−length o￡ ParameciuM．
2） Effect of the medittm temperature on cyclosis．
The velocity of cyclosis was investigated by changing−the tempera−
ture． The medium concentrati’on of artificial sea water WaS equiV alent’
to O．Ol M NaCl and its pH was’ 7．3． The time of adaptatioA ’was’20
hours． The observation was started 5 minutes after’
狽??@tempetature
was changed． The Pavamecintm was observed at higher tempetatuice
丘rstly，．and， after the obser▽ation， a目ower temperature， was again ob−
serVed at the same temperatu．re with the first observation． And ’ 狽?
d・t・w6re e血Pエ6y・d・nly wh6n th・v・1・・ity・f・y・1・si・atエ・st・．・b・P・Ya−
tion was equal as the first vaiue． The result is indicated in fig． 2． ”
hln
this figure， the velocity of cy’ モ撃盾唐奄刀@is indicated on the ordinate and ehe
teMperature on the absissa of the graph respectively， and each cirlr”］e
P1”
’represents the mean value． ：1
As is apParent in fig． 2． the velecity of cyclosis changes linearly with
the’ モ?≠獅№?@of the medi一 i i，’
um temperature， and if
the velocity is indicated
by Vpt／s， and the temper一 B
ature is by tO C， the ’rela一 b
tioh between the veloci一 i／’
z
ty and the temperature is
represented by the follow−
ing fo］rmula， i． e． V一一at十b，’
where a＝：O．1， b＝＝O．7，
be−
tween 50 C andi 30e c．
by
The velocity−change
the
temperature−change in
medium was
reversible in
The
nユost Pavamecium．
velocity
relation of the
o
O ・s． lo ・ls ’ co 2s 30． ec
Temperat”re ’
Fig．2 Effect of the medium temperature’ on
cy，closis．
The velecity of cyclesis is indicated． on ordinate
and the mediurn temperatur’e on absissa． Each
circle represents the mean value．
with the temperature
be工ow sO ’C was expeCted ・veriy i’nteresting， bu．t it
of a suitable apparatus．
could not investigated from lack
3） Effect of the concentration of me
dium on cyclosis． ’1’
affected greatly
It was’expected that the velOeity of cyclosis must be
by the concentration of the external medium， for the Velocity of cyclbsis’
is affected by beth of the change
ef motive force and the change df
the viscosity of protoplasm， and then’ the viscosity−of protoplasm changes
with the change o￡ the colloidai state’of−protoplasm，・ and・the colloldal
52（4）
A． KOENUMA
Nq． 4
state’of’protoplasm is’affected’by the’osmotic pressure of．the medium．
Hence， an investigation Qf the relation between the velocity and the
cQncentration of the medium was carried out．
Paramecja were adapted’ for 20 hou．．rs in the test solutions． Variott．s
concentrations of artificia工sea water were Used， their pH being Set
at 7．3 by adding NaHCO，， and ．their temperature at 269 C． ． The result
is indicated in fig． 3． The ordinate indicates the velocity and thq absissa
the concentratien of medium． The c．oncentration of・the medium is
shown as the mol−concentrati．on equ．ivalent with・NaC］ ・solu．tion．． And
each circle in this figure represents thg mean value of the velocity of
cyclosis．
In this figure， t，he velocity
4
decreases slowly in both sides
of 0．002 Moj； and in the
3
エnedium concentrated more
盈8冠﹀
than． O．02 Mol， the curve drops
2
in considerable rapidity， and
at last in the medium con−
1
centrated ’more than O．1 IMol
the．cycl◎sis disapPears cQ皿一
o
O・001 〔LOI e．工M．
pletely．
It js・sai・d that the osmotic
OonCentration
Fig．3 Effect of medium concentratiQn on
P「essu．「e of p】℃toplas皿of Papt−
cyclosis．
amecium is almost equal to that
The ordinate indieates the velocity and the
o￡ O．05・Mol or O．！ Mol Nacl
absissa the rnedium cencentration． The
medium concentratjon is shown as the moJ−
concentration equiva1ent with NaCl． And
solution． lf this is true， ．it
can．be said that・the cyclosis of
each circle represents the mean value of
Pcdptamecium disg：ppears in the
cyclosis．
hypertonic extern． al solution， in
．． This
hypotopic enternal solutions
other word， cyclosis appears only in the
fact suggests that cyclosis of Paramecinm requ．ires the， entrance of water
into．protoplasm from external med．ium． The requirement．of entrance of
water， into protoplasm． from．the，vacbole or extemal medium in the
protoplasmic ．streaming of Nitella has been discussed by OSTERHeUT
in 1952． lt is very interesing with．the case of the protoplasmic stream−
ing connected that the’ cyclosis of Paptamecium appears in the hypotonic
solutions in which the ep−trance of water into protoplasm・ may take place．
4） Effect of NTa一， K一，’ ・Mg一 and Ca−lons on cyc］osis．
It has ’been well known that various ions in which K一， Na一， Mg一 and
Ca−ions are contained， affect protoplasm． The effect that these ions
given to the cyclosis when Paramecinm is k：ept in the mediu皿containing
K， Na， Mg er Ca ion in varjous concentrations was investigated．’ The
effect of each ion was observed by making the solution in which two．ions
STUDY ON CYCLOSIS OF PARAMECEUM （1）
No． 4
（ 5 ）53
were contained in various ratios． Every test soJntion had a constant pff，
71 or 7．2， and a constant concentration， equivaient to O．Ol M NaCl−so−
Iution． The t’ ?高垂?窒≠狽浮窒?@at observation was 200 C，・and the time of
adaptation of Pammecium was between 18 and 20 hours． Paramecium was
stopped by a narcotizing method where the viscous medium was used
with isopropyl alcohol vapour． Every observation was started after 30
minutes from narcotizing treatment．
The resu］ts are dem6nstrated in the following tables． Table 2， table
3， table 4， table 5 and table 6 indicate the resu！ts in the series of NaCl−
KCI mixtures， NaC1−CaCI2 mixtures， NaC1−MgC工2 mixtures，』KCI−CaCエ2
mixtures and KCI−MgCI2mixtures respectively．
Table 2
VelocitY of cyclosis in NaCi−KCImixtures．
velocity
varエance
一一
O ＃f’s
。D
S
ρ◎4
畿3． り09臼−
o
一 ﹁
5
e．e30
1．5
一一︸
012345。○
1．or
e．612
1．8
0．C・925
2．5
0．075
2．9
C．055
2．7
0．104
3．Q
C・ ． G4・5
2．9
0．e23
3．0
0．033
2．9
e’． ・04．5
3．1
0．025
3．2
0．es2
． Table 3
Velocity of cyclesis in NaCl−CaC12mixtures．
leg ［Na’）／（Ca”）
velocity
／
りO
∩O
C e G
0 0 47 8
4畠
0 0 80
Oり
2 7
0 0忽
O O
nO
7
5 ︵O ︵D 3 4
2 7μ S
2 2 2 2 2
0 02 O
0 0720
∩0
︵D
0 0 4
0 062
﹁
2 5
nO
，
R210123αっ
一 ︻一 ︸
αD
varlance
number of
observations
5
01
010
9．0
0
9
G
31
910
15
11
10
10
1
10
10
1O
1
1
10
6
1 1
1000
log CNa’）／（K’）
number of
observations
No． 4
A． KOENUMA
54（ 6 ）
Table 4
Velocity of cyclosis in NaCl−MgC12mixtures．
／
0 0625
︵0
ら0
・
OD
0 046
0 0
0 0 22
0 01
，
︵0
■
nO
0 07 9
，
几D
，
0、
0 04
0 07 1
0 7
0 0 27
0 02 0
9
．
，
00
． ，
ロ ロ コ ロ コ コ コ ロ ロ の ロ ロ
S
μ
の
T
4
3 2
1 0
．
［
一
1
αり2．345
一一一一
oっ
varlance
019
0 976
Table 5
Velocity of’ モ凾モ撃盾唐奄刀@in KCI−CaC12mixtures．
Io9 〔K’〕／〔Ca層つ
velocity
・
0 0 22 5
，
只り
0 01 5
01
0 75
OO
王
∩0
コ ロ ロ
∩0
02 5
ロ
0 0 1 75
60
︵0 ∩0
0 0 1 00
0 01
0 01 5
0 0 1 00
0 0 26 0
0
4。っ．
． 圃E ． ，’● ． ， ．．
000
S
：．、．﹄． ﹁ ．．
．﹁ ．， ， μ ／
・ 田 ●
47
S32ー0
ゴ1’2
33
00
層㎜．
一﹄
一﹃．︸一
。。
varlance
の
Table 6
Velocity of cyclosis in KCI−MgC12mixtures．
log CK’）／（Mg”）
nD
0 07
06 0
， ．
2 Qり8 り0
00
﹁ ，
3、43 9臼
0 ∩︶ 1 1
0 0
﹁
0．
S
varlance
﹂
0 0 0
買∪
民︾
0 0 0
0 068
0 065
9
nO
0 01
0 0 67
∩0
ロ ロ コ ひ ロ の ロ ロ コ コ の
T43210﹂．1．233．34。。
μ， ．﹁．湘 ．㌦．﹂﹄．﹂﹄
ノ
．一︻︻一［一
4幽7
。。
velocity
numbeゴOf ド
08
01
0ー
0エ
0ー0
0
．
9
011
0
91
01
01
01
01
0010
6
．
61
019
0ー0
010000
1
1
1
10
10
10
10
10
1 9
1 一0
ユ
19
11
1
1
veloeity
22
38
．23206434．3 7330789．45753
82
72
22
32
42
72
82
92
82
83
22
22
2222．221112
02
02222222ユー100
2
3
log （N4’）／CMg”）
observatiens
nu憩ber of
observations
number of
ebservations
No． 4
STUDY ON CYCLeSIS OF PARAMECIUM （1）
（ 7 ）55
In figu re 4， the results obtained in the series of NaCl−KCX， KCI−CaC12
and K：C1−MgC］2 mixtures were drawn in one．graph． The ve工ocity of
cyclosis is indicated on ordinate， and the va門並es◎f工ogalit㎞of the rati◎
of K−ion concentration to the lenic concentrations of the others are
indicated on absissa of the graph． Each oS the syrnbols represents each
of the results obtained量n愈hese observations， i． e． circ工e represents the
velocity of cyclosis in NaCl−KCI mixtures， cross the velocity in KCi−CaC12
mixtures and triangle the velocity in KCI−IN（［gC12 mixtures． Each valu／e of
the figure represents the mean vaiue of the velocity．
松
If the facts in ’these
e
tables and figure were
noted， it is’obvieus that
3
in the mixtures contain−
the cyclosis disappears．
あ二8︻叫﹀
ing much volume of K−ion
’x
2
x
Moreover， it is clear that
．X
工
three curves−in figure 4
’
show the same tendency
in higher K−ion concen−
tration． ・ ln the other ions，
however， ne remarkable
effect given to cyclosis is
to be seen： ’一 ・
When ’the cyclosis
e
一5 一a 一3 一・2 一1 o 十2 十2 ＋3
moo
Lmg
［Na），CMg）pt［Cin］
Fig． 4 Effect ef Na一， K一， Mg一 and Ca−ibns en
・cyclesis．
The velocity is indicated on ordinate， and the
values of logalithm ef the retio ef K−ion con−
disappeared in the K−ion−
centration to the ionic concentrations of the
rich ’so］utions， it appears
Each of the
others are indicated on absissa．
normally agai’n vv’hen Na
ion is added’to the so−
lutioris， except the time
十4
o｛］
symbels represents each of the results obtained
in these ebservatiens； circle represents the
velecity in NaCl−KCI mixtures， cress the
velecity iB KCI−CaC12 and triangle the velocity
af’ter the ’disapPearance is
in KCI−MgC12 mixtures． Each value represents
so long．’ The saMe effect
the颯ea皿value of the ve1⑫city．
is also obtain6d by the additibn of the rest of ions．
Thus， it is clear that
K−iOn iS fev「erSib工e． The CaUSe Of thiS
the disappearance of cYclosis by
be ’attributed ’to such a fact that the
disaPpearahce of cyclosis cannot
coagulation of protoplasm oecurs by K−ion， for the fluidity of pretoplasm
can be recongnized in such a condltionL
If the effect of an external medium on protop工asm is discussed， the
permeability of protoplasm must be considered in each case． According
to AKITA’s data， when Paptamecinm was immersed to the medium ln
which KCI and NaCl were contained ，in various ratio of concentrations，
K−ion content of the cell was always kept over the definite value， and if
the ratio of K−ion to・ Na−ion became larger’ than the definite value K−lon
conten・t of． the cell increased repidly， while Na−ion content of the cell
56（8）
，A．’ KOENUMA
No． 4
．deCreεしsed、reciPro￠ally， becaロ．se an ionic．ex6hange betweeh． Na−ioit in．the
Ce1墨．．a鍛d K・ion． in the medittm ocCu．rred。 The fact that． K：一ion in．externaユ
嫌ed．iu璋was即ore perm：eable．・than．．Narion，．’was．． also．indicated in his
data．：．Thus，、it is：．easy． to consider that a’1arge amouht of K−ion’exchanges
加．Na−iOn．in薇e ce11，．so that．the nature of protoplasm℃hanges． g士eatly；
寛。．．a廷e．ct．．．the、、 cy’closis．、．greatly， whe且the rati（〉（）f K−iOn．concentration to
．Na−io血．．．．concen‡fati◎n』◎f the ．externa1．、．＄o工ution ．is』．1ar．ge．．．But in ．t五e re；
￠iprocaユ．condition，． it is c◎βsi〔ヌered．．that the．cyclosis is．not． affected by the
externa］medium， beOause ， the．nature．of protOplas血does not change so
great工y， for the io且ic ratio of protoplasm∵Uoesロot．國．change、greatユy．
According to HEILBRUNN， the permeabilitY of prQtop1asm toi K−ion oエ
Na−ion is considerably larger than the permeability tσ，．M．90r．’Ca−iOn．．
Hence， it ca皿． be expected that Ca−ion ’oT Mg一三〇n．．in th．e external solutiQh
is less ready to ente士into the protoPlasm than Na一え。ロ， so that the ionic
change．． i且external medium by the fortner ion圭s．工ess ready to．．lintroduce
the． change of protoPlasm． ． 1．、 P．．．・
・／Another fact in the figure and the tables is that、inぬe． solution
c6htaining CaC］2， the cyclosis shows theユ（）壷ef velocity than in the other
Sblu．tio且s． This phenomenon seems to occu．r by a、．decrease of per阻eability
、qf．．・pltPtoplas血， beC．ause the de￠rease of permeability己of．ectoplasm occurs
by Ca−ion in the solutiGn． In the result of the observatioロof．the ve工ocity
of cy’closis in a．・series of．．NaC｝CaC工2 m圭x；tures， an u．nexpeqted． result、that
the cyclosis gets moreてapidity in Ca・free solution than the others， was
obtained．．’But this resu工t may’be explained by the e任ect． of Ca−ion o且
the p晦eabillty・that Ca−i・P sets the．e￠t・plasm．in a less．permeable c・n−
di瞬The・1・gk・f・Ca−i・n in・xt・圃皿・di・m increa・e・．th・p・・me・bility
of．ectopla＄m．of Pavamecium， but．．Na−io且． in．extemaユmedium is kept still
almost． impermeable）hence the．nature of protoplasln do＄s not． change
except the entfance of water increaSes， ahd here the increase of、 the
velo6ity・ pf￠yclo6is．Occu；s． 亙f a．己．Considerable amount of K−io翠i串．con−
tai血6d．｛血’出e extem・乳・り1・ti・n，． th・エ・・k・f Ca−i・n i・the s・1・ti・・、． b・i・g・
ach即ge of．th6 natuエe．of protoplasm， for the K−ion of the sqlution
ex・h・P9・・wit馳・i・nS in th・． o・・t・P1・・m・．Thu・ゆe cy・1・・i・di・apP・q・§
ih the Ca−lack． soldtion in which a considerable amGunt of K−ion is
60ntained． ‘ ． ． 、
In c工osing this paPer，亙wish to thank Prdf．．YUIT正RO IKEDA， aロd
P・・f・HARUq KINOSgTA・f T・ky・Uni…sity・ f．・雌・i・g・id3nce・and
enco肩iragements throughou．t．this work ttt ．．．、tt
s聰搬醗譲y
The ・effects of some factors on cyclosis of Paramecium was Qbserved；
，r．1］， hQsg． fa．ctQrs． w． ere t−he body−length’ o￡ Paramecium， the temperature o・f
No． 4一
STUDY ON CYCLOSIS OF PARAMECIUM （1）
（ 9 ）57
the test solution， the concentration of the external medium and the Na一，
K一， Ca一， and Mg−ion in the external medium．
In Paptamecium， of which body−length is between 160pt and 210／t， the
velocity of cyclosis was not concerned with the body−length．
The velocity change of cyclosis occurred in proportion to that of the
temperature of test solution as far as the temperature was between 50C
and 300C． These changes of the velocity by temperature−changes were
reversible in most cases．
The cyclosis disappeared in the hypertonic medium， and it appeared
in the hypotonic medium． This fact suggests that the cyclosis requires
the entrance of water into protoplasm． ，
The increase of 1〈一ion and the simultaneous decrease ef the other
cations in the external medium made the cyclosis disappear． This dis−
appearance of cyclosis was generally reversible， and it recovered by
the increase of Na一， Ca一 or Mg−ion． The effect of the change of the rest
of the cations was almost negligibユe．王n the explanation of the actions
o￡ these cations in the external medium， lt will be considered how the
permeability of protoplasm plays a r61e to the ions and how the per−
meablility of protoplasm is changed by the ions．
亘』量意er盆重題re
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BILLS， C． E． Biol． Bull． Vol． 42： p．7， 1922．
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HOSel， T． J． Fac． Sci． Tokyo lmp． Univ． Sec． 4． Zool． Vel． 4： p． 299 1937・
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OSTERHOUT， W． J． V． 」． Gen・ Physiel． Vol． 3s：p．519， 1952・
WALLENGREN， H． Zeits． allg． Physiol． Bd． 1， s． 67， 1902・