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The haptonema

Article · January 1994

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Masanobu Kawachi
National Institute for Environmental Studies
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4
. Thehaptonenla
T
5AO I NOUYEand. ¥ 'A5ANOBU KAW
l ACHI
f
mli
ll
l(
l'ザBi
山'.
(
ir
J u
lSc
ini
fl
'，¥， ['川即日 i
t
yoJT
5'
l11
1
110
{
/，
jゆ (ll

Abstra
ct

.
¥1
，orpl
1OIogi仁川 ，l xhav i
oll
ral，andph ysiologicalf cλtuJ't:
s0'1Lhehap l
0 1
1em< l<t!
'e
rcvi
cwcdandt hei
'!biological i，
gnifi仁alcei sd i以 1川町1. Tht 、l川ptoncr n礼 l
al
臼 )
11
1
1川 nen lぷ a r
γ λn礼 Igedi l
l川 1abs ollllCc onfigllration inr cl
at
ioJ】 to 【he
fagel川 c
l lpP<I
l l
'，lUSancl仁ι・
1I11
1arorgani .a
'
l ti
on，andt hed ireclionり fI
wndi ng
anc
l coil
ing i sal s
of ixed. Coili
ng i sa Ca 2--I
ll
t'd
: iλled p ll(.
:
nυrnenon.Thc
adhesi
veprop erLyofl heh apl01t汁I川 ISP日 ) babl yrcl礼l cdl Ol hep resenc eor
.
cnnc丸山 町 Jii
l
l九b i
llι
lingsubs l
anccsand/orel ectricchar 貯 九ssn仁ial
'dWi
t
: lhl l
ie
h出plOnemal alsur f
ace
.Hapwnem alalf uncl i
川 15川 epro vis
ionallypropo sed

Introduc
tio
n

Thchap WlC l llaisat hrcan- l

ikt' or ganelel halsuper fi
ci al
lyr cs
cr nhlesr he
nagel l
u1ll.Thi suniqucυr ganclcwasknownas・l hel h
irclr la
gcl lur
n・lIn ti1
Par keI 'la.l( J 955)r ecogn izじdi tashci ngc 1istinctf r01ll r
lagelaは l l
tlgι!V(' i
t
rhcnamぞ ' h<lp lOnema' .lLhassof arI Jccnl uundonl yinhapl ophyl es.
Thcfundanwnralprop er ti
esofl hehaplolじlIIawcrcr cco gni7ed i nl he
perioc lf rOlll 1950sL o car ly 1970smai nIy by nri lish phycol ogists( f
or
bihliography，s eeHibb引 d 1980) .Thcyf OlLnnr h<
lt:( 1)The孔xonじr nc0 "
[
lhc haptoncm< lcOl llpris
ess ever alsi ngl ct Illicrotubl l
le s，t :
ypicaly目I Xor
seven， wh iιh arcs urr ollndec lbyaci SlerU<l0' 1r hccnr lopl<lsmi cr eL
Icu lul1I
(ER )川 I dt hcpl aS
Il ll
l< ell
lll1a;( 2)ThehaploncmaG.1I1 < ldh eresl Os uiLablc
SlU
L Slrataand丹howgl i
ding l lloliliLy;( 3)l ns or ncs pc仁 icst he hap lonem辻
仁oi I
ss ponl <lne ouslyυr whcn phy si
caly s l il
llulll
< ecl，aldυ1 1 chcll1ical
fixaliυ1
1amlccldcat h;( 1
')ThehaplonじH I抗 仁訪日 bcnd，b l
lti ldoe snolbeal
l
ikc a nagel l
llll1.Al ll he昌c f c川 ur cswcr c 仁 omprehensi vefy1 じvicwcd by
Hibb引 d (1980).AlIlong t he manye< lrlya rticlcs pu blis hcd r he f olow-
11
1日 比「じ 【h cn】ostimport ant .Mal 】1υI (1 <J6 4，1< J(8)i nvcsligat ec
llhe u lLra-
stru<"tl
lreofL hehapL υICIJI辻 inc1ct ai
l<Incll .eac lbealer(J971a)i nvcs r
iga red

ThfHI
f/
lI/
(
!J1
ザIi
:，¥
，
l
IJ山 '(
ed.J
.C.Gre elぬnds.S.C.1
. 1
，・1h
c c
;1IC
'r).S Y
Sl'J
t nat
ic
s: ¥
.ss
uc
i 泊 。 nSp c
cj九l
¥
'Oll
ll
lH
、う)
.
- 0
..i1.1 '.7
'1 3-8リ C
I‘t
rendonPn剖 . Ox
for
r
l.1 9~H . iJ)Th<:oS
ysl
<:
OlIa
T ti
csAss oc
iali
oJ1.
1994
74 Th
l'Hゆ 1
0/
11
1)
' "r
11 l
lgnl
'

lhebehavi ouroflheh< lptol1t>lll

a lI ~ ill g 仁 111 ふ III i<
.:ro
s c
opy.¥;[司jorclc~crip­
l
iolsoft hi
sor g<ll1el
leappωri ng il1p r cst'lll -dayt Cldbooksofp l1
¥'col og ¥'
are bas c
cl0 11l hcscpi OlH'ぐ nn g wυrks .l I o¥ ¥'
e¥ ' er.il
刊で s r
igal
io mのn I h is
org日I1cl<、hav cmadcs lυWp lυ認r essd l lJ'i ngl hep< lSIr¥
¥'
CI1I
)')
'('
ars
.
Int hisc1laplcr，wt :wi
. lJ'C町iewIllorphol o gical ，h cltavυ
ilIral
.a l
ld ph ~rs io­
log
ica lf はl
じ
， UJ'esυfl his unique or g日nelι i ncor poratil1g (Ll
la reccnl ly
υbtainccli n Ol l
rl aboralOTプ( Kaw<!cl tial1d Inυ1I)E' 1 094: ha wachif'lnf.
199り .

Morp
hυJ
ogy

111 、
1泊 1 membe rsofl hePr Yll1lc sial(・s.I l
tcl t a pl υnCll1ai swe lldc¥ '
cl opcci
and f Ul1uiυnal .l hali s，iti scapahl 。
、< rc oilil1gandac lhe s
i ol 1 0CXI (
'rna l
垣uusl rata.1 Li spani cll arlylongand CO I I昌pl仁 UOU ~ i ns pecies01 ・I h(
'gCl1us
Cltつ ・sor !
JI'
OI ItI Jf ina，c. g
.I Ipt o1 60μ 川 ￨ υngi nC .m 1/ld!n(Lcaclbc<lI CI・1 971( /
)
(Fig s4 .I H，4 . 26). Thc l ong an c lf l
lnCl i onalt Ia p l υ l1el l
la，a lso，i swi c
it>l
y
c
iis lrihlllcdi lll hecoccol ilhopho ricis
.
Thenumberofmicr ol llbulcsi ~ fix仁df orapa rti clIlars pc仁1 '昌. bUll
.
( her c
are¥ '
aria li onsberw (、(
'n叶)('仁 i cs.ThcuS lIa lnUll1h ( 、ri ss ix()J'scven( Fig.4 .1 )，
bUI i n an l l lld(s
、 crib じd h aplophy tc wぞ ha ¥ 'c1 '( '(
'(
:'11t')discovcred，t hc
hapL OI1C'lll ac o nt ail1sf υ
，unc c・nmicr Ol llb
l ll .s(
p ドi g.4 .3).Theul traslrllClll代 、

ofI hchapwllcmal all ipha sbet >1l im '

(・slI galedbyManLOn(1 964，1 96~). lt
isnOl'll1alynal -endec l，b llll apcl'cd.J .
l υi Jlled01 's pa lll
lal (‘fortl 、
l昌 弘 lυOCCllr
(
ルl 迅IIIυn 1 064) .Manlon Lll ld Lcむ dal e ( 1 96~) and Lじ泊 dbcaler (1 971 a)
ιl
ell10nS lrat ed thcI H'
(
'S Cllceu[aswel l
i ng r haLl l
ll 川 ピ supa ncl dOWIl 11<'
haplonema." 1・hc1 11り rplりυ l氏i c
alba日i s0' 11 hisdy"泊 I ll
icS lr
llcl 1r
l ehasIlot
beeni n¥ 'csl ig辻 l CU.
11I ltcI r じep artoft hehapt olcl l
la，l lres ha ft.m icrolll hlll
csarcar ranged
I
Is Ia
l lyi nar ingS l
lrrolll l
dcd U) 'l hrccconct >n lr i仁 川 じ l llbrancs( F
ig .ι ι1 )，
ll
reinncrI WOofwltidl泊 1 ・eofERt h日ti >'
!(' ontj Il Ll u lI ~ wilht hepcri pltcr alER
υft hcccl .
l' '1
1 1
(・I r辻ド1υIIer nal alERal so見I rυw saC- s h日pぞc iprohki nL l司ans-
¥'
er se~(， c l i o ll ， whcr ci ti sl inkccl1 0l l
rc pl a sl1la mCll1hr al
l仁 b >' bridges'
(¥l
1 anL OI1 1 964 )
.Tl w c) 'lnplasmiccor <
:仁uJll ai nsr iho s orncs.bUll l
1(:
"1
・t'h as
bccn 1 10 J't: ponofan~' 01 hcr仁o lls li
lllenl Smakingupl l
re haptol( 'mat a
l
sh<t[t. Hのwever ，in1 11泊1) '品pe ci e
s ，日 ne !( 、
nrolldcns es tructllrci 同 pn
:
， scl l
la l
[
11(' C('l1lrcυrl l
licr otub lllarr il
lg，む υI llecl ings i ngl ctl l
liuり lubulc九 loonc

Figs4 .1-4. 8 ( 1)T ranwlT引~t' ('1 I

OIIo(ahapl onCllIa(りIllpr I
sIng~C \'C I1 ~Î n 只lel
mIrr<llll
hl
llesSUlTOl
ll
lclcdb '
}I hrヒヒJllt:
ll1branc s(Chl)'mrh川 IIIlIfi
llιh i
lla)( x11 (
)( JO (J)
(2)F .
lt
-nl'OlId enses tr
uClIJlT C川 11
1円 '1IJl符JllIc
r Olubul阿 川 1 (1dt:ns edOI~ ( αI T0 7
llC
)
SI
ll
lalcd( )
111 heり UL
erI 1C
l l1b
l ranc01 '11(' I'
:R( /
)IYllme
silllJ1!
H II7II
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()).( 3)
Hapt Oll
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l (
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l‘ 中引・ H
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lIngf o unc l'
ld円 t nll
ldens cIllI C
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-
l
llbuh'.ぉ (
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lalM吋 l ionofaha pt ol
ll 'Jla~h()町Îng l
l hcbaql
 .( 1コ
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，
lS
;
) I;
S. )
A I
SUC.よ (
8).
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cscq ;
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)uuO
コ(mo.l.w) s.n.~q ;}SU;}([ ( ~ ) .
(0 f
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7llUtOA1/JOfUl/コ); L
lqll;
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76 TheHa
pto
Phy
teAl
gae

anOl h er( Fig.4 .2 )( seeCregson e ta .1

f 993) .Simi la rS l rucl urescan be
[oundi nl hehapl onemal ao[Chrysucl ‘υ'Omulina chi ton .( F i
g.3i n~an ton
1967)anc lC .αthele s(F ig 22i
目 nMo es trupan clThoms en 1986).J nt he
spa cebe tweenr hぞ hapt onematalmemhr anean rlt heour ermemhrancof
lheER，t he rea res omes lruct ll
reswhi 仁 ha ppeari nr rans vcr scs cctions泊S
dul Ss ilU辻1 じdu nl hcuulcrERIfICmUr allc(F ig.4 .2)0 ['de!ls ebar sconne Cl-
ingt heouterERmembr aneanc lt hep la smamembr 九n e( Fig.4 .!'l
)
.
Bef or een tering i ntot hεcel，t he micr otubul csar c or g辻 nizcd i l】 泊
trough- l
ikear rangcmen ls llrrolJnnc dhyar ingofER ，s!owing辻 c
l -orarc-
li
kear rangemen rinr rans vcrsf's ccr i
o n( Hihb cr d] 980) .Thじ 仁 on f
igur alion
01'l hc何 T m仁 川tIlh l
l1csi nr c
lar iol to. thcf lag ellarapparal usandcel lular
organC'1 1csi shxcdsot ha t(01 ")oncab su 1ulcc onfi gur alioni spr e
senl.The
(
'00仁川 Fじ s i dco[l h
cmic rul ub u1詰rl J
'ough[ acesoneo[l hef lagelat hati s
refer redL Oa slhel e[lfla gelum( Inouyean clP iena ar1 9H!'
l) .Thcmicnト
lllbul arc on figllration a ts ucc essi vever ti仁a1l cvclsofl hc hap l
onc l
l la is
show ni nF ig.4 .1 H.
11c l oscproximi lYl Ol h eba se，l hehaplon emai sof tens w olle n( Fig.4. 4)，
llJOug'h swel l
ing i s abs ent i ns ome s pec ies， e. g.P ηmnesium pa11Jum
.
(YIanlon1 964 )
.Thec yl opl asmoft hisr e gioni smor cc1 cctr ondcnsel han
int hcd is la1r egi onandt hemic ror u
b lllar亡。 n自glr alIonl
. a kesl he[ormo[
ana rcorha lfc i
r cle( Figs4 .n，and4 .1 H).ThcERi scons ic
ler a b1yr eclucecl
andf r agmen redi ntotwopor tiuls :onep υ r
l ionl iesa longt heconv exs i
rle
ofl hぞ ml 亡rotllb1l1辻I 羽 仁 川 l dl heol heral ong t heoppo si
r cs i
dcofl hc
hapr oncm凶 ( Fi
g.4 .6).A5f ara swehavei nves ti
gar erl，t hcb出 a lswcli ng'is
nlosl仁 ol ls picLlOUSi nCl nyso chrom 叫η iataγkeal!( Figs4 . 7，4.8) .
The[ reepa rto[t hehap tonema，s l
igh llyd is.alL
t Oil Senl lγi nlot hec el、
ismorpholo gicalyu nic 1l
J c( Hihbcrd1 980) .TheERf ormsaf ingcr -likc
proce sswi 1hat l
alcnd cXl ending L O 、
¥i a
rclst he conc avc s idc ofl he
microwbul ara r c(
‘
Fi氏s49
u
目 ，4 .16 )
.Thecy ropl asmi(仁or ci sJlloJ'ee1 ectron
d(、ns cl h泊1 l batalanyot herl evels ot hal ，il
ll ongi ludinals e ct
ions ，i tis
SCCIl 泊S迅 l ransver ses eptumconnc仁 t ingl hef ialend oft hef inger ・likc
proce ssandERr nembr aneofr hcOppO Si
l Cs ide( Fi広.4 .J6). Thi sr cgionof
lhehaplonemai s.caledt het r a
ns i
lioT! r egion・Thehapt onemai sof t
cn
det achedar ound. thisIcv clWI ICIl i
li sdi scar c
ledi nr espons ctophy si
仁川 υ r

Fig
s4. 9-4. 17 ( 9)T r'a
ns vl:!rses e cl
ion01 'thctransi
tionr e符ion01'th eh apL Onema01 '
Pη川 IfSIUJIl/1(/1(1
1
11.s
1
1 l
tow in gt het ing e
r-ilk
cp roccssofER(刑判川) ( x1 11:(り0
) ).(10)
Transvcrscs cc tiυno f'!
heha p.lonemao [Cllt
yso
ch r
omul inaalif
em 川L11t" l
evel ofenu γ
inl
uL hec el lNo
. tct hca rc - l
ike; ;
I
1'
l'<
llgeme11l01
'm ic r o
t u
bulcsasinF i
g s4.fiand4. 9
(xI 1(
)りO (
) ).( 1 1-15 )Tr ans v
cr se町 nion s01
'theha ptOne maa tSuccl's
sivel !v
t !l
t s1'
1
'01
11
dist
;
t1! opr oxima l( Clt
l，'
s
! o chlOlIlulinah ir(
f
l)
.出>I.t! I
_he仁on l
igura t
iona lc hang c(lnt!
aci
diti
o lof日 目、，mi cr u
Ol bule s( numb crcdi古川ld9) ーl !n
Jt
: sema t
erial ar
isingf romt hc
 7
ilehaやl
one
m乱 7
7

16

ちるJ
ccntr coft hcm i cr otllb
lll a ra r ci sSCCIl in 1Lλnd 1 2
.Nl Imbcr in
gi ， af
tcr; ¥
:1
0c写tru
p
andTholl1s cl( J 9京6 )( x16000) .(16)Lon gIludinaLSCC
tiOIl ofthc【r日n s.
t
ionr
i cgi
on
(Clp)'soc
hr01 l
ltltin作 s p
.).Notcf i ngcト l
ikcpr oc
c ssofER(m γ'ow) andt hc n -
ans同 rs
c
sep
tu lll (x 5(000). ( 17) Lon符i LUdinal s cc
tiol
l of a c oiJ
cd haptoncm 羽

(C/ny¥l)d ml1lwlillfL a l
i戸川l ) ， l)i rt>n ionり rcoi li
ng istowardst l 、
w con' exs idt
>0 1t
' he
Ili
Cf'OlllJl
I"
l ，
r 11・('， An刊行， illd i c a l 円 IIH~ l
il試刊 帽l
ikl，
'(
>ro
('
('sOrI・~ R (
x410(0) .
7
8 TheI
Iai
やlf
J
oh:
ヲl
eAl
gae

chemi calsti
mllli.Ma mon( 1964)s ug gesledl hallhisspe cialr egiりn辻仁 l s
ω
a‘lock ingdev ice'o[u leextendedor ganele .
Thehap tonemaen tersthecelatade fin
itepo sit
ioni nr el a
lio nlol he
nγof Ia
- gela( seeGreena nclHori，Chap t
er3 ).Mi cr o
tub ule sext en ddeep
intot hc仁' )
tosols howi ngco凶 gur at
i
.onalchang es(F i
gs4. 10-4.1f>)wh ich
hav ebee n desc ribedi ndelai
li ne a r
lyp llblicationso[Manlon ( 1964)，
Le adbeat erandManton( 1969)， andr ecentl'b
) )
'Moe slr upandThomsen
(198 6). 加 exa mpl eo[ahaptonem atalba se，t h
ato[Cl ny sochr omulinah i
rla，
isgiveni nFigs4 .1 1-4.15，andr ep resenteddiagrammatica lyi nF ig.4. 18
.
Twomi crotl
lbule sar ea ddec
loneb '
)onei ns equenc er eslllt
i nginn ineal
themostproxima Iend.Thel1li crotubularar cbecome sr earranged t o
show achara cterisicpro行l
t e，us ual)'r eferredt oa saz ig-zag( F
ig s4. 1
1，
4.12). This compr i
se st wo supcrpo scd 辻r cscach madc up off our
mic rolllbules( F i
g. 4. 1
4). Imm edi ale
l y above l he le velofl he d islal
conne ctingf ibrea ssoci a
tedwitht hebas a
lbodie s，e l
ectrondens em a
te rial
ispr esenta tt hec entr eoft hel1lic
r otubularar c( Figs41目 1，4 .12).Th is
l
1lateria
lext endso vcr辻 ndbcyondt hcconnect i
ngf ibre( no lshown) .I n
themos tp roxil1la
lr cgionnincm icrotu bulcsa rear rangedi nsuc haw a)'
thate veη'けl reea dja仁c ntmicrotu bulcsform迅ncqu ila
ter all ri
angl e，and
scvcn ofl he
s e ar e he xagonall
y packed ( Fig. 4.15). The haptonema

A B

e を
立 s

f
。
。
00000

g 。
。
o
00000

¥ L
¥ d ~ / R
h :;0
。。
o~~

今
年
Fig
s4.1 8D iagramma icr
t cpr
cscn
rari
onoftheS l
nlC.
lur
e0"1aIypic alh aptonema.
(A)Tra ns
ver
sesect
ionsofahaptoncmaarsucccs
si
vel e
ve1sfromdis l~l l 10 p
roximal
cnd.(s)J心 I1gi
s l
ud川 alse
ctj
onofahapt omena( proxi ma
lpart);(泊)ー ( i
)in( 八)
cor
rc s
pondto( a)
ー(i)in(日) し=ler
tf
1agell
u m:
R=r i
ghtf1ag
cll1m.1¥ ・ 1mb
l cdngり f
microt
llbl
ll
esaft
erMo出 L r
upandThoms en(J986
). Nottoscalc
.
 Theh
apt
one
rna 79

lerminat esnea rt hcl eflb asalbo d y.Atv ar

iousv erticall e velst hehapto -
nemai sl inked1 0bOlhba s albod iesby5 CVCI出If ibr ousban cls .A fibrous
hap tonema talr oolar isi n
g[ r ol11the1 1
10stproxima lenc lo[t hehaptonema
andd i spcrsing
-i ns i
clet hecel，i sknownon lyi nl heP a vloval es(Gr eenand
Hihh erd1977 ;Green198 0)
Thel eng【hoft hehaptoncmav ar
iesatdi f
feren ts tag c
sofc clc yc
le.It
shor tensdurin gr cp l
icationo[t heflagela rbaptonemal
・ alcomplexs oUlat
itisv crys hυrli 1c elst halha v cjustωrnplel e
clceld ivi
si on ( Ma ntona nd
Lccd迅l c1 963;InouyeandP ic naar1 988 ).Thehaptonemai nιrc出 csi l
S
Icngl hi ns ever alhou rsa ft
crc cld ivisi
o n.Regener at ionoft hcbapton ema
wass ll
lc
liedexperimcnt aly(Lea c
lbeater1 971α;Gr egs one ta.11993) .AIter
exci sion bybubblinga ir，l hehaptonemar egener at cd andr ecoveredi t
s
110r mal lengthi n12-1 5h.
ThchaplOnemai srudimen taryorcv cnc n
ti r
elya bse nli nt heI so
ch r
γ-
sid辻 l
e sands omeco c.
c
.olithophori ds.lmantonia' ro
tu
. η
d α，Di crat e
r iainomata ，
ancl mot iJe. celsoft m
・ ilian iahu .xlのi lack haptonema t辻1r nicrolubules
(Kla v cncss1 972;Gr een an c l Pi enaar1 977).Othcrs pec ies ha vea [ ew
11
11仁 rOlu bulesi nt he shafr;r h rcc 川 Ch ηs ot
ilal arnello
s aand C .s ti
pitat
a
(Gree na ndPa rke 1 974，El75)andf ivei nJ s ocηs
h i
sg a
l ba na (Horianc l
Gr een 1 991).Somes p<河:I
(川 υ [p必，u.mc hワs isanc lHymeno 仰1 ηω ha
0 ves hort.
andbul bollshap: toncma tac o n tainingupt os ixmicr otu bule ss urround cd
byi r
r egularly泊r r辻n gedER( Man tonanc lPet erfi1 96 9;Lcadbcat er1971伐
lnouy ca ndP iena ar1 985).Thehaptonemai sus u
a llyrudiment aryint he
Pav lov泊l e
sbei nga t tenua tedandcon rainingonl yoncυ tf ou rmi c rolubules，
lhoughr ht' nnmh e rinc reasc si n日id cUlCc elt os evenorei g
'h t( eε.Green
1 976;Greεna ndHihb c
rd1 977; Gayralandf resnel 19 79).

Behaviourandph
ysi
olo
gy

A
.dl
tωo
io
nandt
rans
loc
ati
ono
lex
ter
nal
tar
tir
的

Thel taplonemai sc apableofcoi li

ng，bending，and( )
fadhcsiont ocxternal
subsu'ata The
目 s
e heha vioural f ca
tllrcs ha vc uccn demols trated by
Leadbea ter( 1 971α)wi thex:ccllcl
lt【 :
ine- p
ho. ographs
t .How ever，thebi o-
logic
aIandf ln仁 l
ionals ign
ificanc eoft he scr novcmcnt sha slongbccna
puzzle.Ther unctionalsigni
fi仁an仁coft headheringn at
u r
coft hchapto -
ncrn凶 W 辻 Srecenlyc lemons lraled byT <awa .
chiptαL (1 991) 1 nr hemixo
ー -
lrophics peci es(;fnysor
!n
，
J οmuliη(l l
urta，thchap t onem誌 pl
aysaprima ryrole
infood仁 抗 p lllre.Pr eypa rt
ic
l esa c
lher et ot hehap ronemawh i
lstthccel
swim swil hl hehaptonemapr 吋c cting込hcadandwi thlhcflagelab eat
i ng
辻long s
id cl hccelbody( [
orwar c
lsw imming )( Fig
.4.19)
.TheC ldher
i ng
parti
clesa r
et rans l
o仁arclc
c 10w n. 0apar
1 【i
仁 1I1
辻 rpointonU' I
Chaptonem辻
約
() T
hl'11
αII
[oj
Jh
ylf'Algnf'

Figs4.1 9
-4 .2
4( 19 -23)High 叩 e edv ide oi ll1ag-e~ ofpre ¥captlr'i
t lO l l)' . wr h mm u l 川 (l

11
111
0.( 19)Adhe si onandt ra nsln印 t iOl1of1p ，
' It
l i
cka ll dformationo faggreg a t ea t
tt
Il'p'
lIticll'礼ggrc g al
Ing(Cl1trc(P AC)( x1 200 ).( 20，21)lJp¥ v
ardtl 司1l ~ IO C;'lti c. ll l 01
al1aggr e符a lefroll1 thePAC tol ht:' i
lp( x1 60() ).( 22，23)Dc li
vCI下o [t hca g g r e g al c
tot hcC l'lIsu r
fac cbyb cndin宮 o[l heha p lollema( x1 6(0).A円 行1/IS ild( i川 ぞ t tll'
dircctiono[t ran s1ocai
lonofpani cl's
t .( 24)Ct'l lllt'ndi
llg
-tIλptoll'llIa¥ 'i t ho lll
capturt'dp ar
tiιl
e( x1o)
(O)
.

試IJ
υU L2μ md isLalfr om t he ha se，caledt hepar tideaggr ぞgating(ent r e
(PAC).andaccu ll
lulaL cL h<.
:
rcr <
.:
su lL
ingi l
lt hじ dじ目、 l opmen toj 'ama sミl
Y{'

aggr egat e( F
igs4 .19.4 . 20). ~vIor pholog i ca ll )" ， the PA 仁 pr obab lyC O
I・I・
ト

spondslO t hcbas alswel l

in g.I nt heaggr ('
gat e .indi vi
dtlalpanicl e stigh tly
ad lt Clじ lOりIlじ辻1 l0ll
tcrs uggcsL it
lgl hats り]fH
.'仁C l l
lCtltlla L
cn凶1i ss('仁lTt <
、c
ta l
thePAC.Ol lce. υnned，t
[ I
Je社符符 rcgal c 1l10VCS1Ipwardsa1 1【11・ 凶 仁h 山 l
ltc
hapLOlemal altip (Fig s4.2 0.4 .21 ).Theh ap lυnemabend si ntυaS i
gT 10i
l d
shapesoona [
tert hea 日r
定 egal ear r
i vesall het ip(Fi g.4 .22)孔nde ¥'enlualy
de l i でr
、 si
tr ot hecels urfacea tt hepost eri o renc loft hecelwher ei li s
in g est cd into al oorl¥ '<!nlolc( ドig.4. '2
:1).I kn c
ting ( )fI he haproncma
occurscveni ncelst ha ldonott rans locatea~~rc~a tcs ( )fprc)'( Fig.4 .'24).
l ，
Li wo川 1ωtill只 L haLL hcωI I仁C出 c s!IagclarbC<ll ilgjus Lbdorc 1 tc
1
ha pwn('mas L
ans[ 0hend.an clr hatr hcf lagclas hows udrlcnandvi gor ous
ac ti v
ity when t he aggregat ei sdet achec l[ r om t he hapronema. The se
phClOl nCl
l辻 suggCSLL hathapl OnCl l
l込laluchav i ouri spr c仁isc
l yrcgu l辻l cdi l
an i ntegrared C unct ionals ystem，r he n 乳g ell ; lr-hapwnemar alcompl (
'x
Cl oscf l
lnni onala ssociationbcrwccnt hcha pt oncmaandt 1<::
.g
c
llal1lay辻 l so
υcClIrduringt hepr ocesso[c oi ling(~ee be low).
The di rertion ofhenc t
ing dlr ing aggregat ct rans portt ol hc c (
'1
以 l r facei ~ f
ixedi nr el a
liυII1 υcel lularorgan iz 孔lion， Thi 5wascun fir med
u s i n~ Ch ηsoch1'Omulinastini(em.anoL hcrs pcci c sl halllscsLltchapt oncma
 Th
eIwt
lonema 81

f i g.4.2 ι4. :34 ( 25)Vi dt"oim< tg >o

t fC lu) '
sοrhr01l lulinl7l tinijトm， showin 符t wouncf J
l.
lal
Il agcl‘ 1andcapLllr cdmic ros pbc rc s01t hcbapl OIll'lIla( x~OOO ) . ( 26-28)仁o llf ocd
l a~ t' r mi cl'ogr aphofCluysοchr ol1!叫 JU/s t山斤r a.F Iω r csccnccofFI TC- Iゆ elc d(o n-
( 山 川 日l ilA i s1 りe<1i
t 庁 dり I llhぞ h泊]11 り1It'f1λ (
1 26)I' h 出ぜ仁川 II 川 ~ I ; ( 27 )引 I ]
)ellll
lドり s e
d
mi crngraph01pha町 C Oll ll'
司sl;"tlぞ1f 1uol "f，c
' cl)cri tg
ll1;
" rs;( 28)f 1l
1on 、"cC'nc('111 tg
;
: C'
.

( x~.j()()). ( 29)( ;，川 .

f "od
l !10J Iu
l liu(/1 t
i11 l Wi
1 lhl hehapl onemat igh. l
lyc oiled.S p ine
s c al es日r ea lsos eel】 ( x1 800) .( 30， 31)¥ 'ic
leoima ges0' 1C ;hr:;s
oc hroJIIlll
inah iγl
a:( 30)
ct'1I capl l1rin再 baCl eriaw il hhapl olwma :( 31)celW il hal l lidi
n e-coal cdl ar
exmi cro
喝

、 plwr t" < 11 <

1<I wd 1 0l wc
l t
'1Ih od')ands pin('s c司l es( x1 2(0) .( 3 2) CI υ)'
.
I"o
(l
'!rmnululIl
h irl( /("("， 11 n，
:;)1
. 円 1 ¥
¥
'ilhp r o(λl ilt'.Tht' 1 1;1]
)1日 1
1
("
'1)1 1 I
1
- 可t'X1 t
'1Id (
'(1.1)111 partlyc o川
Ai
1( ，
:dλ
吋
刊
， 1
川1
l
t hct ip (x1 800) .( 33， 34)Dι c吐1
l辻c h礼
l'l 泊pl
dh
叶 υHι I
日
ぐ寸
引
. 1
1
川礼 0
日
川 1
"、(:h
η
川?)
'叩
Sο
EJd
.円

恥1'1
日l hproc ω
ainc( 33)an 】CIf olow cdb
吃 η)'A231 87( β34
引).NOL Ci ncluc tionofcoi l
in g( x1 800 ).
( トigs1. 32~J3〆1 f rol1l Kaw 礼ι hiandl nou' )t'1Y9' [)
82 Th
eHαp
top
;
lり!
t
e11
lga
e

asafoo dca pl ur ingd ev ice.Thi sspeci e sha sl WOunequaJf lagelawh ich

canbeus ed a spo sitiona lmark ers( Fi g.4 .25) .Thehaplonemaal ways
hendspa ssing betw een t hetwof lagela，c losert ot hel ongerf lagelum
( Fig
.4 .39A).Thel ongcrf 1agel lu l
flc orres pondsl ot hel e flf I迅gel lulll/
bas albodyandt he[ oodv acuo1 ei sl oc at edsomec 1i sl
ancebel owi l( Fig .
39 A).Theco nca v es ic
1eoftrougbーl ikemicr ot u bul es[ ac est hel eftba s a l
body and t he d ir ection o[ hapt onema t al bendi ng i s t her e-
fo rei nterp reted a sapproximat elytowar dt heopen s id e0 1' thet rough
(Fi g
.4. 39A)
Ther εi sno r lo uht.t ha tt h ehapt onemat a
ls u rfa ccmc mbranci sad y -
nami cs tr
ucl u re.IIow ever ，i
ti spr esellL1}7unknow nwhatkindofmechar ト

ismsa rei nvo1 ve c

1i nl headh esionandt ra nsloc ationofpr eypar ticlesan rl
whatkindofmol ecul esmigh tber espons ihlef ort hc scphcnomcna.Thc
adhe si vena tureof. thef lagclars ur fa仁ch 誌s bcc ns wdi edi nChlamydomonas 目

Ch lamydomon α scelsa ltacht os o lidors emi solids u r[ac e

sbymeansoft hei r
n迅gclaandt h ewhoJ ec ellca nmoveonl hes ub s trate.Thisphenomenon
isr ef erredl Oa sgJ idingmot i1ityorf 1agel 1a rs urf acemot il
it y.slood g oor l
(1 990)reviewedf 1ageJ la rs u1'facemol ilityandl is tedf Juest ion品 t h辻 ts hou l d
beans wer ec
li nf ut ures tudie s.The f olowi ng q u cstio n s mayal so be
appl ica blet ot h ehaptonema:( 1)Whath叩 LOnemaL alsur [ acecomponen ts
ar er espons ihlef o r.themcchanic alf or cer equi re clt oa c1he1'et oext er nal
pa rticlesorot hじrsub str ale?( 2)川 市 ati sl he1 1
1 0t o1' 1
110lecu ler espons ibl e
fo rgcner alin g movemento[ext erna lp ar ticle s?( 3) Howi s【hch泊 p LO -
nemal a1s ur[ acemot ilitys y
st em r egu lat εd? ( 4) Wh ydoespreyp articl e
adhe si oninduc et ranslocat i
oni ns tca dofcoi lingo[t hehaptonema ?
InCh的作りdomonas ，ilha sbe en1'evea led【ha tgl ycop rot ei nsa rci nvolv ed
inadhcs i
onofl hef lagela( B 1o oc lgoocl1 990 ).[ ti spo ssi b lcl hats imilar
sub stancesar ei lvo
1 lve c
li nhapt onema t ala dhc sionands ur fa cemot il
it y
.I n
orclerl Oe xaminet hispo ss i
bilit y，weaddcdFI Tιl abeledconcanav al inA
(C oA) t hats peci flcalJy bi nclsα・0・gl u cosじ andα-D-ma nn osei ntot he
medium.I ti si n tere sting t hat，i ll
I hec a seofChη! sochror nu linas Pinザe ra ，
f
1uor e scencewasdct c仁 t
. cclon1 ')υn l hehap tonema ( F
igs4 . 26- 4.28).I t
ShOl ll dbeno tc clt halno f 1uor e sιencewasohser ved on t hcf lagelaor
C.s f
Ji ηiJaa ，wh ile Chlam .
wimno ηιsf lagcla s howed b righ l nuo re scen仁f
undcrl hes amecon cliliols . Re s ultssof arob t aincι 1i lldic a lel h司 1 lhe
hapl onema t
a lmemhr ancc onta insgl yιυpl'Ol ei ns01' gl ycol ipi dst harbind
wi thCoA，a nt ll h 以 tl山 me mbr anei sd i引 i ncrf rom t hcp la smamel1lb rane
O[bot ht hef lagelaandc cl bo tly .Thiss ugges t
st hatt heha pt oncr nat al
l1e
1 l
11h rancf orr ns凶 t l
is tinc lmembr anedomai nonChrysochromulinac els.
AnOl herpos sible f o rce t ha t仁ol lldgcner辻 l eadhe sion i san el ectri仁
cha rgea sso仁 ia tedwi lhl hehap l onema t almemhranc.V Vci nv cslig泊 te
ι1l h is
po ssibilit)
'us ingel ectrica ly仁 h ar 又c dl alexr nicr ospher es
.Whenwcaddcd
mic ro spher eswi thanc g at i
vc仁 haq{ e(仁a rbox y l
ar eoralhumin -coatcd)i nlO
 F
heh
α 0ηe
戸1 ma 83

the CU IL
Ure of C h rysochmmulina hu -
ta，microsphcrcs a lla
ched L Ot he
haptonemai mmecliatel y.A similare fl
cctwasal sof oundwhenb act
e ria
wer e use
cl( F ig.4 .30)
. The c el surface ofbact criai sb elievedt obe
ne g
ativel
yc har ged.I nιont r
ast，p a
rticlc
swi 山 apo sitivecharge，s ucha s
ami dinc-
<:oatedmi crospheres，a r
tachedt os pines ca
lesr hatcovert hcι ell
bod y but not t ot hc haptoncrna ( Fig
.4 .3 1
). Th iss u
gg'est
st ha【【he
haptonemar als ur
fa ccofC h ir
lαhasapos i
t
iv echarge， anc
lagai nsupp orts
thcs uggc日tiont h辻tthehaptonema talmembranef ormsad istin
ctdomai n.
¥，
¥， けl
ヤ crcforcp os
tu latelhalanel ectri
calcha rgema ybe， a tleastpar tly
，

1
'cspon邑iblef o1'adhesion.I tisgener al
lybelievedt hatt hecelmembranc
isn egative
ly chargec ls ot hati fO l
lr int erpreta
t io ni sιor r
c仁 t
，t hc
hapt onemat almembrane i svc ryunus ua.
l Morc c lct ai
lec
l physiological
s.u
t dicsarcnccdcdt osubsLamiaLet h
ishy poUlesis
.

C
oil
ing
Theseconnpromi ncntf c a
tur cりfh辻p LoncrJl辻t は1bchaviouri sc oil ingand
un仁oi ling.l ft hch辻 ptonemai swelde velopedandr elaLiv
e lylong， coi ling
IS込 山 liver salfeaL ure( Fi g.4 .2 9)‘ al lhoughsomes peci e
sc oi
lt hei rhapt o
-
nemaLa1 1
101'e re adilyt han o ther s( Leaclbeat er1 9 7 1α). Manton ( 1964，
19 68) putf o"叫 r da' wr gorhypothe si
s'a sapos si
hlcmc仁 han isr
nf ort.hc
coiling ann uncoi ling 0' 1hapt .oncm a t
a . Shc p ost
ll l迅t cdt ha L仁oi li
ngi s
inrluc cr
l hy辻 仁 h辻ng じ o ft ll
r g'
or pr cssurci 1 t
l hc ERcavi ty， 辻I1d t h a t辻
gra dua linc reaseofi nL ernalpr es su reb uilLupL oprodlcelInc oilingoft he
haptonema， whereasaq u ickc 1ecreaser esultsi ncoi ling.The reha v ebeen
nod atas Ip
l porrin gt hisf a月cina ti n gideaand，a tpre s ent，rher ear enod ata
Locx pl ainhowr nc仁h辻rI1仁 a lf or仁じ sinducc仁o il i
ngandlIIlc oil ing.
Coi li
ngi sfrequen tlyo bserv eclwhencelsar eswimmingr api dl y，w ith
nagela bea t
ing behind t he cel ( backward s wimming )
. Manys pe ci es
undergoforwardSWi 111
111川 gasano rcli
narγmodeofI OC01l10t ionwi tht he
h泊 pronem辻 p rojcctingahcad.1 11s uchs p巳ci es ，rapidba ckw ar ds\o\~ m 111 ing
O仁仁 urswhcn仁cl仁ol lidcwi th ob st
.a仁Ics
.On cont a ct， cclsi nsta nt lyc oi
l
lheirhapLOnemaLa， unc l ergυas pinningr novcr ncnt，r
. cvcrsct hcr l irc仁t.i on
and i ncrea set he s peed oft he nagelar heaL . Ce lss wim away f r om
obst aclesi nt his way ，impl yi nεthatcoi l
ing pr oduc e s an i m mec li aLc
出 o ir
lingr cspons c.lri spos wl at ed，t herefore，t hatoneoft hefunct ionsof
thehaptonemai sobsl acles ens ing，all easLi n日peci e見 Lhalus ualys w irn
forwardw it hthehaptone1l1司 di rect ec
lahea cl( seeals oGr egs one lω .
l19 9 3).
Wh ens p
υ lItalleouslyl rig只ereJ， coi li ngis出仁仁omp迅 nicdby仁u rlingoft he
f
lagela ( Leac l
beat er 1 971α )，s lggest ing c lo sef unct io nala ss o cial ion
b<.
.
:
LwCCIl thchaptoncrna辻ndHagcla .
Coi ling i s an eXl r e1l1elyr api d pr ocess， l aking 1/100s t o 1 /6 0s
(Lじ泊 dhca tc r1 971α ).1 ti scven l ess rhan 1 /200swhen condi tionsa rぞ

favolrahl e( llnpllヲlis hedobs er vations )

. When compared W iLh coi li ng，
日4 '
j'
Iz，/
( /
(/j
Jlo
!J
h_
)'
(ri
¥(gae

F
igs4 .35-4泊 ( 35-37)Sequcnlia
lim.lgcso{c oi li
ngi n Chりsoch
r iIIl1 1
o/1lllf mJ71lI
:
¥01'
(I halc o
ilingslar
l"川 、r、
't'r
alI
川 I
1
11、 (x~(JI)O ). (
3 8)Scanl】i
ngIlli cro只r apho f
(
;1m WrlUOIIIu
/i110s
p・.l
hchapwncmac oilcdi1ac
l lo c，i
k
¥ 、
ehelix(x~.ï 0(0) .

In
l coiling i samuchs lowerproce ssandlI~lIally l akcsS C¥'cr辻 1s ecυnds.I l
begins[romt hebaseandp ro
c ('( 'dsIOwardsl hcl ip
.Sincel hehaptonema
ぞの i l ~ whcnc eIsan
.l 、 仁h el
llic辻1)'白xc d01' dl ie cしi lha sbeens lIg gest ed1 hal
('
oi ling i sl hc 1m んじl
， e r向 。語la t 、
e (Elepancl l '
vlac lnr)Te 19 Í'l~)) 日nd l hl
lS
lll lωi liIg i !
>l heener 即'-c
lepcndent[ "
01'111. 1 i spo ssib lcL o make haplo-
1e
I matar epeatc oi
li ng日ndlIncoi lin日bvr cpc泊 L ingphy sicals t
i ml lli，s lcha s
lapping11(> mic ro s仁opc.) -Iowcvcr ，r じ
いe< 11ecls hockreslllL~ i ng rad l
l all o
s s
0 1 ・ ~ens iti vit y (l
.caducal cr1 971ω ) .Cυil in g!> la rL~ mO S lh で<lll(、
I 1Ll¥'fr oll
lL hc
r
lis talt ipandpro仁ccd邑Lowardl heba、 、 bUli
巴 lm爪.al sob egil1 a l!>c¥ヒr al
pO il1l Sa l olgl hじ I cngl h( Fi良s4 .35-，1 .
37)
Thed i rt :cuunofcoi l
ingi sfixedwi thr ( 'sp cclL OI】じ 仁 un fi試l lr ali onoft he
hapLoIlemalalmicr ot ubule sandc cllorgaJli L込l iυn.Oe l ailec
lobs er¥'ationof
coi linε¥¥'as mad. e hy Lcadbcalcl ( 1971 a) l Ising high-~pecrl ci lふ
mic ros co p y.Hcf o ul ldl hλ1t hピ hどI i
xwasl or med0 1
1l hcs idじ t υwar d!
>l he
cc ll .i .e
.whel ハ iewedl r oml hc，¥( '
1 1Irals idc.t hchじl ixi >
!1 υrmedm r hekf l
si deoft hehapwllcma (ドI 符・4 . 3 9B，C ).Thehel ixofr hch辻 pLOlc lllaI >
!
lo r l llcdi ld p r ecis epau削 ' n.O ll rohs cry ati ols hU¥¥ lh alc 10ckwi s ehe lic('s
ar eal waysrormed( Figs4 .3H..~. :39). Tlted ir ect ionofcoi lil1gi sthusoppo ・
sil cto l h cbじ ndingl ha loccur sd llring p lで)' l r
all旨pり I し i.e.，lO、~'a r clS111('

u
υm' exs i c
le0' 1r hemi crolllhula r.1I仁 υ fl hじ haplonema( Fig.;.3
1 4A).1 1
1l l
tむ
h( ， )ix，t h cr nic
r ulubuksareλrrangf'dt o' 101'111< 11a l
仁 W il hilSconc川 ' es ide
faci n gtowardst h(
'九xi s01山 じItじ l iA，whじr e asm i crO lu
h lll csa rca rr<ll 符cdi l
l
ac ircl
ι:whenl hehaptonemai メi n< 1nc XLcldeuc υnd il ion(Gr egson1 1n
' .
/.
E)9 ~3) ，
 The}
u中t
one
rna 8
5

Fig.4.39υia grammat icr ep

re sent
ationo fhaptOnemat ashowing山 ed irection
so f
coiling川 l db ellι
li
l
lg! -
Olt hetr
"anspor
l01 "a g
gl"
eg 孔L
e討 (A)Cln yso
dtr
o川 u{
i咋() c
elwi l
h
al1leriorflagelaandhaptOnem a.Co ilingisL O
w ardsU1Cco nvexsidcoft hcm icro・
lUl ml"l辻 氏 。 r lhelraplOl1ema， andbel1d ingisL OWλrdslheoppn s i
Leside.( B，C)
Ven tr alandlat eralvie
wsofC I!lyso
chrom!tl
i押a cc
l
lW ilhlatcr
allyinsc
r t
cdflag c
llaand
hap. onem
l a.C o ili
ngoccur sloward日I
.hed orsa
lsideo flhec elandi ntoac lockwi
se
Ir
el.ix
.F v=loo dv 泊仁lIo
1e;1 1= }川 pl
one Hlt;し = l
< etf
f l<
tgell
ur n;R=r i
ghlt1辻g dlurn
.:
¥0 1tOs
. cal
e

Si n仁じ bolhfoodιapturingandcoi li
ng凶 r ca sso仁i atcdw ith p'1n icul'1r
movemen loff iagellaasmenL ionedear l
ie r， ilisp osllllatedl h凶 lh迅 plonc -
lll
atalbeha viouri sa f fect edbyt heCa~- concen tration i nt hehaplon e-
rnatallumcn出 F ort hcchang t
‘o ffl'1gela rbe a tp a
tter n01'otherorgan isms .
Tυ l ωl l his hy pOlhcs is，品 目ic r辻I e x pe rill
le nrs wer e des igned us i n
g
oり sochromulinαhi r
'l“( K awachianul n ollyじ 1 99 4).Wc白r stusedt heCa 2+
ionophor e A231 87，which gene ra tcs 仁h辻nnclsi nt he cell memhr 1n
' e
throughwh ichCa~+ i onsc anpa s s.Sincel heCa 2>con仁cntr 辻t ioni nsidet he
cel i snor ma llykeptl owerthan t hatol l
l side，t heoccurrencc of5 uch
仁hanncl sprohablyc aus esani nfluxo1 'Ca 2+ionsもVhenA231 87wasaddcd
lol heI nediulIl，coilingwω induc cdi nl l
lostc els( Fig.4. 2
9 )
.I nanot her
expe rimen t， we added EGT A t ot he mediuIll l o rcdu仁c t he C'12 -
仁onccn tr川 i on.1 ns uch a con dil ion，coi li
ng wa ss uppr essed and L he
hapt oncmaw'1skep tun仁ol iec
le ven i fc elswer ephy si
calyS limulal edby
tapp ingorc hemic alf ix alion.ThcselwOcxp cr irncntsi ndicatct h a
tcoi ling
isaC泊2 +
-mer l
i'1tedphenomenon，a n cli sc ausec lbyt heinDuxofCa2 +from
oUlsidel hec ell
.Simi larexper il
llcut swcrclInder t
.akenbyGreg sone la l
(19Y;))u引 ngEGTAa n cll anthanumch lor ide， ac alciumc hannelb lo c
k cr
.
Th eye Slimatedl hall hellue~ h り ld 仁υ n 仁じ ntr辻 tion ofc xtracellu
larC a~+ i ons
forhaptonel llal
al仁 川 I ingt o0仁仁 urlie sbeれ、! een 1 .
O-7M a ndl O-6M.
8
6 ThrHa
tJ
1oj
Jh
)'
trA1
9ar

ILha sbeen[ Ollndl hal ，il

lllIus dcc cls ，thcERi sus ccla saC日2 -poolan cl
Ca 2+i onsr el四 月e df rom i tinduc emus cula rconl racLIon ( I
illu1 989).Thc
ER. Ihats u rr ol.lnclshap lonel l
la tal ll
licr otubu lesmay ，the re[or e‘beanO Lher
po ssi blcs ou r仁じり fCa 2 +supply.[ ti sal soknow nr ha tc af feineinducesCa 2七

inc1uced - Ca -
>
2 r clじ出む(じ l CR)f rom. t1
、l 五RpoolofIllus
< decels ，anclt hat
pr ocai nei nh ib iLSi L(Iino1 9 89) .vVhclIc affcincw泌 出I dじdt ot hemcd ill
ll
弔l
問0 9 {o[celsco iled l hei r hapLOIlcma la.1 1
1c onu ' ωL，t hc 仁 oi lingwω
sup pr es se c
lb ya clcli n宮p rocai nel Ol hemed illl
ll.I IapL Oncmal ac 1id1I0L( oil
ar3 1 仁川 町a~ecl coi lingatan i nt ermec liat es t ages ot hatma nycelswi l
h
p迅 ruy仁o ilじι Ihaptoncmat awc
. r eohs crv ccl( Fig.4 .:12) .The seohser vations
sugge sll hall heER i sa lsoi nvo lv cd i nl hc仁o ilingI IIC仁 han isll1a ndll1a y
sen 'ea saCa 2~ r e ser voir.When celspr e-t
real ed bypr ocai ne ( sol hat
coi l
i ng 01 't he hapt onema tawasi n hib ited byi nter rupt ion o[L heCa2 -
surr1 y)WCI T cxpo sedt oA2: 11 R7r heh九p tonem証t 礼 c oile c
l.P robab lyCa2
ム

ioni nt luxfromt hcou tsidel l

led il1ll1 incll1ceclcoi ling.F igur es4. 33and
4.3 4a rcS L
l仁 ha ncx ampl e()fdcr a chc dhapt .oncmat .a.
I
ti sobv ioLlS[ rol11 lheaboveexperimen lsu lall h ehap lonemai sah ighly
Ca2 '- c
1epenc len l01 又a nele .Perh apst hec ha ngeo[c lirectionanc lbeat ing
frequenc y0' 1f lagelai np η' mne si ophy tecelsi sa[ s oduet ot hci nfl1x0
l 1
Ca2 +i onsi nt ot he f lagelaan clt hei rha sa lr cg i
o n日， a sr hisi soncoft hc
univ cr s alf cat u r csoff lagclaofr n証nyf lag clla
l cor g辻n ismsandga IllじL
icc c
lls
.
Wi lhl he s eυbs erv孔lionsi l
llJIi nd， weL h in kc oilingi sl h ef trstphenomenon
o[a na vo iclingr espons e.Gr egsone ta L( 19 93)r ea仁hedt hesame亡 。n chls i
on.
Themechani calf or ccwh i仁 hgcncr atc s仁oi lil
lg抗 nd/orb亡n dingo[l he
h7lpr OIlf:'I1l71 i
spn、s cn rlylIn kllown Nり dynei n-li
k eS lr uct ur ei sa ssociatec
1
、
会

wi thr ni cr otul Ju lcs.GI でg sone tal .(υ1993) haede 引11

10J1幻
句St
叩r'aぽ
te
， ds幻tハ
ruκct
印ur
吋e s
Eh
出 川at(
. ωo】n
町r 川 仁1 泊4 辻仁 C川 h 泊叩
}ド
判
}孔L
υω
I口l I
口
児亡引ma
叩 t
凶a
討1mi 叫icr
比 ol
凶u b叶
u.l
1 est o oneanot h
山河e
、 ran
sugg じ呂 ledl 山halas 凶l
1idingmec ha n is m ma ybei nvolvedi ncoi ling.

Co
ndudingr
emarks
Thc rci snodoubll hallhehap lonemai sa1 110reco mplexor ganclci n
bOl hl llorphologicalandphy si
ol ogi仁alaspc 仁tstha n b泊spr じv ious l
ybeen
Lhoughし Taki ng int oac counrohs ervar
ionspr cv i
ous lymadc by many
phycol ogi sl
Sand t ho seweha vema de，wepr ovisi
onaly5 1ggeS Ih
. apl
.o・
ner nal alf unct
ionsa sf olow5:( 1)Thchap. toncmai sas ensOI)
'de vicef or
der ecti
. ngobs .
tacl
csa ndpOl CIlL
i辻 lyuns l
lilableext ernalenv i
rolment s，ancl
coiling i spar to[an avoi d
i ng respons et hatgcncr at c
sI 辻pidb辻仁 kward
swimming ;( 2)1 tisus cd a sade' ice[ 、
01' caplu r
ing，aggr egar ing，anc l
Lransponingpr ey
.Thea dhe sivepr operlyal s omakcsi tpo ss
iblef ，rcelsl
υ O
alta仁 hw 込 ne xl
ern社 1Sl
lbSL
r ale
.The res eell
lr oheall e
ast[¥¥10 dif
lセr cnt
pr o印 耳 目est hatforml hcph ysiolog i
calb asisoft hesef unctions，i.e.aCa2 +
-
mcdi at cdr csponscandh泊 pwnem ar
als urf ace11l0ti
li
ry.Alf lnni onss eem
 Thl'I
t
r
.中I
OTll'lla R
i

Fig.4.40 D iag
-rammaucr cprc sell
lal
io l
lofv ario usphcnor ncl las ufur !仁o
f g
'nized
inthehapt oner nぬ (ω'xn linsol
i dl i
ne)a
s ndt hcpo ssi
blephy " iologicalc vcnt s(boxl'd
indolll'
flI川 内) thatalowl hcs"f
t u
・llclions
.I n O
l lxofCa2+ i Ollsf !
"ol
llnu .
tsideo rby
calc
ill1l-
l i
ndu( (
'd- c
alcium- rel
easc(CI CR)f rOIll lw 1 ・
:{ma
J )'
ind llcccoi li
ng.Agg r t
!ga-
tionofpr c
)'pan i仁ksI Ilayher ealizedb ylhcS CCl"Cl
jOl
lofct-'lIlell
lrnal erialar oun d
1heh asalswel l
ing.COlC<llIa 山 l
il
lA-bi ndings ubS lancc s01'ckCl ricぬ1cha r gesonr hc
h泊 p
tol
l ell
lalalslIrfacemaybcr cspυIs ib
leI (I' I
) hea dl】esionufC XLCIl1<1
1P出 L icks' "
toas ubs tra
tc.:¥1υ1りl'rnolecule
sl hatgcnc ral cs ur[泊仁じ III
Oli
lily川 Hl l hemechan ical
orCt"Sf
f orben rlinga rcpr c
scll
lly1 I1kll
<川 11. Suw./lnr roo
l
lsindi cal(l
， hcd ir e
Clio
ll( )
f
sIr
l faccmotiJ it
v.: '
¥otlり おcal
e.

lu bep rccisel
yr eg ulated i n<ln i l1
1 cglはl cdf l
agellar・haptonematals 戸tern.
Po ssib
leh< lpl0川、 rnat .
alfunuiol昌 ar cs urnmarizedi nF i.4.
g 40.
Thc haplυIICU la i san or g<
lne le IIll
iC
JlIC 10l he Ilaplophy la and no
hornol o只OUSs l
rllctur e haseverh(，(，1I o!J品目 vcdi n otherorgani sms. I
ti s
lhcrel'orcυrcOls i dera ble inlelでs t10 know ‘ Wltal i!
>l he origin ofr he
haptonemu'Ollr1I11th.:rs La
lldillg01 t hisorganelei s，howcvcr .still誌L礼 1
iniL
ials lageandwehavenoinf ormati
. ol lO辻llswcrl hisquest ion (butse e
Cavalier 只mirh，Ch辻 ptcr22).Mωlyp
- l obl
・ emsr ern<
lin1 0 hesol vcr
lbcforc
wes arLlO u
.
t i以;u出 l heo riginoft heh< lptoncm辻 al l diL
品じV υI UL
Ionaryand
phyl ogener ic signilic辻 n
- c c
. 1 1IvcsLIg
'a
ti υI1S into the phy siological and
l
lIolecu larmechan is msofI hef1agcl lar.
・haplυne lll
alalcomplexmaybevel γ
ll
11port泊lItillLisr
h c
::spe Cl.
88 T
hl'JI
a少loj
J ，
h)u'i
¥l
gaf
'

Rer
ercnces

l oodgooo，R.八.(
占l 19 90) .Gl i
di ngm ot ilityandf 1ag-ell rg
a -1 '
}co prOI (:indy n川 1 1ICSI n
C h lall lyr
! o
lll(川町、 In C泊。 ' " andJ
) l agelt oTI I
Il'mbraneJ， ( ed.R . A.Kloodgood)，
pp.91 -1 28 .P len ll
llll 'r t
:ss，¥ ewv
' .
川 k.
Estcp，K.W.an c
lM a cll ll yre，ド (1 ~~間~). T泊 xo nom) '，lif cc yc le.c l
i s
l r i bul Iol1a nd
da Sl1lolrophyofCh) ， '
s orhr OlIl
1rl
. iJ/u:" l heor y司ι仁り unt ingf ors cal cs
.hapLOnel1la .
1 I1 1("ifemu
" sbo d icsandt oxicil Y.Marinfルυ1 0J9
と
， 1l hrrr 日九〉川町 5 7，11 -21 .
Gar ral ，l' 川 dFI 円 nel ， J 目 (1979). ど ~anlhe川 adll)'sis 局内:y mlÙlf Lepa ill(~ IH' ( 'r
I '
) llllws io-
phy ceae Pav
守 l
o¥'ales):11 1出 I r uc tur ee td iscus siり 1t axinomi qu e.Pr oti llο
/ο炉 、a，15，
271 -8 2.
Cl t'e n，J .c .(1976).:¥otcs011thcnag -el larappar alu s日ndl axollo lf lyり fPav/ol'{(
/f11'¥
"(l
I)'
rhn 川 、 官 nd
( er¥ -欠 :er ，a nd01 t hes tat u sofP(wlo引() Bl lc
I herトmc lr el込 l e d
g-f- n(~ ra wil hint heI la ptりp hyce日c. .
f O(l円 wl0/I heM{(r in
l'Bi o l
o giralA .
¥ wr ial;(/ (ο
/ Ill r p
[7 n;I('
(1Kin gdο/， 56
1
( ，f
} 9!，
>-fi()2.
Creen.. J
.C.( J980) .Thel ineS Ir
llCl meofI' f lvlovatinguisGn :むna ndap l -
climinal '
Y
ちI I r
"VC }'o ft h cord Cl'Pa 、-
'lo¥'aJe s( 1
'1γmnesiophy cea e) .-/、
山
h I tl' I
t)'m f (/ gimlf.o/(I"II{(/.
1 5，1 51 -91 .
Cret'l l，J.C .and l Ii bb erd，D.J .( 19 77 ).Thc u lu羽 l ru c tur eand l a xonomyof
IJ ，I品削f
刊
、， 叫IWII
叫仰イ
，
U}
'
，，ロi1I
h川;H
日 川/t
{
川
口?1 (ド'
削 YH
】t
川日Ine s
引1りp hy c
山 む泊附c 、
、
) W.V川h s pcci a
川1 r
凶 "
伺cfcr 巴n 可c e1 ωot h c
haplωonemaandf la広e larapp m
】a
加加I削 I 15戸
附 H
川υ
t
川"
円川
、
'I
I(/{1ザ
，
ゲρ
.
'[υ
j
1 附 /1
.1(
，
川
J
けγ
?1
、
γ
川 n p品 川智i mlA s¥or ialiollo f
t h e【J niledKin gdo 問， 5 7， 1 1 25-3 6.
Creen，J . C. <ITl d Parke，M. ( 1974 ).A r c
- i nvc st
i gati on by l ight and c lc CL r o n
llIi cr os copyorHullllem ! 、 lP
.fI(
" /bil川 Ce ille
r( 1Ia] 1t ophyc e 町)，w itlts pc ci allTfer ence
L Ol hei IneS lrUCI IHeor/ .oirls. .
/ 川 In l
a l( // Ihf'! Hnriup8i (/I
(/r
;imlASI(/ ("i(
(I
;(//l o(1 ft!'
United必 ngdom ，54，5 :19-: >0
Grccn. .
J
.C.andParke .r .1.(1 97 5) .Newobs e rv ationsuponmemb erso[t he符enus
C h lyso lilaAna nd，wi thr cma rksupont hc irr el at ions hi p swit hint h eトl ap tophy-
仁e ae.f . ouγn ( /
I 01t heM al 'i nesi olobricalt l
s soc loti o
n o/Ihl' Uniled Killgdom，55，
1 1
) り-' 21
Gre en .. J
.C.an c
lP ienaa r ，R.N.(1Y 77 ).Th et .axonomy0 1t h eor cle r1 5 0 chl γsi clale s
( Pr y rn nes iophι y山 じ) w it hs pじ仁川 1r cfc re n仁C t ot hc g仁川 島r a{ so( hりs isP ar k ι ¥
/) i cml 刊α7 P， I
l'keanι 1I maηI 01/iaRe ynold s.
}<οuma/o/lheルf ar ineHiolo~((l l Aμ o cialI on
ゲ1 1
1(，{ !nitl
'dK il
l gdo 1
ll，5 7，7-1 7
Cr c g son，A .J ..Crccn，J じ. ， a nd Lcadbcat c r，B.S.C . (1 99 3).S l rllcl l lr
c and
phy siol ogy 01 't le 1 川]1l olel l
l川 i lr :
J1I)sochnnnulinfl (I 'r YlIl
I esio p lr 、e
司 ae) 1
ー 1
.
¥ t echa nis mso[haplonemal alc oili ng
-andt her eg ene ra t iol1p roc c ss.f .ou!7wfof
P h y col og y，29，68 6- 700
Hi b b ercl，D.J .(1 980) .P rymne si ophy t es( =Haptoph ytcs).1 11Veve/opment~ i ll II/ar in f
1)1ο lll，
y
;
r ¥ 1 01.' 2，I'hyli柿lK~l/(J l川 ( 引1 .I 乙 R . Cox)，p p . '2 7~ー引 7 ヒI se ¥'ierNor th
ト1 011礼nd，NcwYork.
Huri，T .an clGreen. J じ.( 1 9Y l).Theu llr a5tJ'UCl ur eoft hef 1agel Jarr OOIs y stcm of
hoclu )'
s ;ga
i fbana( ドηt 1nn csiophy la).jOl11la/0 )、tie1
/ 'v1(
lIinel3iofogi f(/ l l \s~ociafio ll 01
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heUnil
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nouyc，1
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of山
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、
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