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Doyle, Alex and Martín-García, Rebeca and Coulton, Arthur T. and Bagley, Steve and Mulvihill,
Daniel P. (2009) Fission yeast Myo51 is a meiotic spindle pole body component with discrete
roles during cell fusion and spore formation. Journal of Cell Science, 122 (24). pp. 4330-4340.
ISSN 0021-9533.
DOI
http://doi.org/10.1242/jcs.055202
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4330
Research Article
Fission yeast Myo51 is a meiotic spindle pole body
component with discrete roles during cell fusion and
spore formation
Alex Doyle1, Rebeca Martín-García1, Arthur T. Coulton1, Steve Bagley2 and Daniel P. Mulvihill1,*
1
School of Biosciences, University of Kent, Canterbury, Kent, CT2 7NJ, UK
Advanced imaging facility, CR-UK Paterson Institute for Cancer Research, Manchester, M20 4BX, UK
2
*Author for correspondence ([email protected])
Journal of Cell Science
Accepted 13 October 2009
Journal of Cell Science 122, 4330-4340 Published by The Company of Biologists 2009
doi:10.1242/jcs.055202
Summary
Class & myosins are dimeric actin-associated motor proteins
that deliver cellular cargoes to discrete cellular locations. Fission
yeast possess two class & myosins, Myo51 and Myo52. Although
Myo52 has been shown to have roles in vacuole distribution,
cytokinesis and cell growth, Myo51 has no as yet discernible
function in the vegetative life cycle. Here, we uncover distinct
functions for this motor protein during mating and meiosis. Not
only does Myo51 transiently localise to a foci at the site of cell
fusion upon conjugation, but overexpression of the Myo51
globular tail also leads to disruption of cell fusion. Upon
completion of meiotic prophase Myo51 localises to the outside
of the spindle pole bodies (SPBs), where it remains until
completion of meiosis II. Association of Myo51 with SPBs is not
Introduction
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+∀ )4%+∃ &41∋%0 1//(5+∀3 )4%− )( 121∋) )( 2+,,%∃%∀) %6)∃1∗%//./1∃
%∀7+∃(∀−%∀)&8 94+& 1∗)+7+): 41& 1 ∗∃.∗+1/ ∃(/% +∀ )4% 2%)%∃−+∀1)+(∀
(, ∗%// ,1)% 2.∃+∀3 −%)1;(1∀ 2%7%/(∋−%∀)0 1∀2 +& 1∀ +−∋/+∗+)
∃%<.+∃%−%∀) ,(∃ )4% ∃%3./1)+(∀ (, ∗%// 2+,,%∃%∀)+1)+(∀8 94%&% &)+−./+
∗1∀ =∃+∀3 1=(.) ∗:)(&>%/%)(∀ ∃%(∃31∀+&1)+(∀ 1∀2 )4%∃%=: 1/)%∃
∗%//./1∃ −(∃∋4(/(3:0 +∀ 122+)+(∀ )( 41/)+∀3 ∗%// ∗:∗/% ∋∃(3∃%&&+(∀
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∗∃.∗+1/ )41) )4% (∃31∀+&1)+(∀ (, )4%&% )5( ∗:)(&>%/%)(∀& +& ∗(∃∃%∗)/:
∗((∃2+∀1)%28
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)4% &.=&%<.%∀) ∗(∀?.31)+(∀ 1∀2 ,.&+(∀ =%)5%%∀ )4% )5( ∗%//&
ΒΧ%)%∃&%∀ %) 1/80 DΕΕΦ1Γ8 Α.&D0 1 &%6.1/≅∗:∗/%≅&∋%∗+,+∗ ,(∃−+∀0 +&
)1∃3%)%2 =: +)& ΑΗΙ 2(−1+∀ )( )4% ∋∃(?%∗)+(∀ )+∋ (, 133/.)+∀1)+∀3
∗%//&0 54%∃% +) +& ∃%<.+∃%2 )( ∋∃(−()% ∗%// 51// 2%3∃121)+(∀ 1) )4%
&+)% (, ∗%// ,.&+(∀ ΒΧ%)%∃&%∀ %) 1/80 DΕΕΦ∗ϑ Χ%)%∃&%∀ %) 1/80 DΕΕΚΓ8
94%∃%0 +) ∗(/(∗1/+&%& 5+)4 ()4%∃ ∃%3./1)(∃& (, 1∗)+∀ 2:∀1−+∗& Β&.∗4
1& ∋∃(,+/+∀ 1∀2 )∃(∋(−:(&+∀Γ )( ∋∃(−()% ∋(/:−%∃+&1)+(∀ (, 1∀ 1∗)+∀≅
dependent upon actin or the septation initiation network (SIN);
however, it is dependent on a stable microtubule cytoskeleton
and the presence of the Cdc2-CyclinB complex. Οe observe a
rapid and dynamic exchange of Myo51 at the SPB during
meiosis I but not meiosis II. Finally, we show that Myo51 has
an important role in regulating spore formation upon
completion of meiosis.
W.∋∋/%−%∀)1∃: −1)%∃+1/ 171+/1=/% (∀/+∀% 1)
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=1&%2 &)∃.∗).∃% )41) +& ∃%<.+∃%2 ,(∃ ∗(∀?.31)+(∀ )( (∗∗.∃ ΒΛ.∃141&4+
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−(7%−%∀) +& 2∃+7%∀ =: ∃1∋+2 ∃%1∃∃1∀3%−%∀)& (, −+∗∃().=./%&0 1∀2
∃%<.+∃%& )4% 1∗)+7+): (, −+∗∃().=./%≅1&&(∗+1)+∀3 ∋∃()%+∀&0 54+∗4
+∀∗/.2% 2:∀%+∀0 2:∀%+∀ /+34) ∗41+∀& 1∀2 )4% ,+&&+(∀ :%1&) 4(−(/(3.%
(, ∋DΚΝ ς/.%20 W&−Σ ΒΞ+>+ %) 1/80 ΜΝΝΜϑ Ψ+∗∗(/+ %) 1/80 ΜΝΝΣϑ
Ζ1−1−()( %) 1/80 DΕΕΕΓ8 Ο∋(∀ ∗(−∋/%)+(∀ (, )4+& %7%∀)0
∗4∃(−(&(−%& 3( )4∃(.34 )5( ∃(.∀2& (, ∗4∃(−(&(−% &%3∃%31)+(∀
ΒΗ131∀ 1∀2 Ζ1∀13+210 DΕΕΜΓ =%,(∃% )4% (∀&%) (, &∋(∃./1)+(∀0 54%∀
&∋(∃% 51//& ,(∃− 1∃(.∀2 %1∗4 (, )4% ,(.∃ 41∋/(+2 ∀.∗/%+ +∀ 1∀ 1∗)+∀≅
2%∋%∀2%∀) ∋∃(∗%&& ΒΧ%)%∃&%∀ %) 1/80 DΕΕΦ=Γ8
Ξ:(&+∀& 1∃% 1∗)+∀≅1&&(∗+1)%2 −()(∃ ∋∃()%+∀& )41) 417% =%%∀
&4(5∀ )( 417% +−∋(∃)1∀) ∃(/%& 2.∃+∀3 )4+& &∋%∗+1/+&%2 ∗%// ∗:∗/%
+∀ =()4 −%)1;(1∀ 1∀2 ,.∀31/ ∗%//&8 !∀ −1−−1/+1∀ )+&&.%&0 −:(&+∀
!! 1∀2 −:(&+∀ [ /(∗1/+&% )( −%+()+∗ −+∗∃().=./%≅(∃31∀+&+∀3
∗%∀)∃%& ΒΞ9∴Ρ&Γ0 54%∃% )4%: 1∃% ∃%<.+∃%2 ,(∃ )4% ∋(&+)+(∀+∀3 1∀2
1&&%−=/: (, )4% −%+()+∗ &∋+∀2/% ΒW∗4.4 1∀2 ]//%∀=%∃30 ΜΝΝΦϑ
⊥%=%∃ %) 1/80 ΜΝΝΣΓ8 !∀ .∀+∗%//./1∃ ,.∀3+0 /(∀3≅)1+/%2 ):∋% !
−:(&+∀& /(∗1/+&% )( )4% &+)% (, ∗(∀?.31)+(∀ 1∀2 )( )4% /%12+∀3 %23%
(, )4% ,(∃%&∋(∃% −%−=∃1∀%0 54%∃% )4%: 417% =%%∀ +−∋/+∗1)%2 +∀
,(∃%&∋(∃% 1&&%−=/: Β!)121∀+ %) 1/80 ΜΝΝ_ϑ 9(:1 %) 1/80 ΜΝΝDΓ8
/)4(.34 ∗/1&& α −:(&+∀& 1∃% +∀7(/7%2 +∀ )∃1∀&∋(∃)+∀3 ∋∃%&∋(∃%
−%−=∃1∀% ∋∃%∗.∃&(∃& )( )4% &+)% (, &∋(∃% ,(∃−1)+(∀ +∀ )4% =.22+∀3
:%1&) 1∀2 ∗(∀?.31)+(∀ ).=% ,(∃−1)+(∀ +∀ )4% ∋/1∀) ∋1)4(3%∀
Usti%&∋o ∗&y,is Β916+& %) 1/80 ΜΝΝ_ϑ ⊥%=%∃ %) 1/80 ΜΝΝΙΓ0 1& :%)
Myosin V and fission yeast meiosis
4331
∀( −%+()+∗ ,.∀∗)+(∀ 41& =%%∀ 1&&+3∀%2 )( %+)4%∃ (, )4% ,+&&+(∀ :%1&)
∗/1&& α −:(&+∀& Ξ:(ΚD (∃ Ξ:(ΚΜ8
Η%∃%0 5% +∀7%&)+31)% )4% /(∗1/+&1)+(∀ 1∀2 ,.∀∗)+(∀ (, )4% ,+&&+(∀
:%1&) −:(&+∀ α Ξ:(ΚD8 /)4(.34 )4+& −(/%∗./1∃ −()(∃ 41& ∀(
2+&∗%∃∀+=/% ,.∀∗)+(∀ 2.∃+∀3 )4% 7%3%)1)+7% ∗%// ∗:∗/%0 5% 417%
2+&∗(7%∃%2 )41) Ξ:(ΚD +& +∀7(/7%2 +∀ )5( 2+&)+∀∗) ∋∃(∗%&&%& 2.∃+∀3
)4% &%6.1/ /+,% ∗:∗/% (, )4+& :%1&)8 ⊥% &4(5 )41) Ξ:(ΚD 1∀2 Ξ:(ΚΜ
=()4 /(∗1/+&% )( )4% &+)% (, ∗%// ,.&+(∀ 2.∃+∀3 ∗(∀?.31)+(∀0 54%∃%
Ξ:(ΚD ,1∗+/+)1)%& ∗%// 51// =∃%1>2(5∀ 1) )4+& /(∗.&8 !∀ 122+)+(∀0 5%
&4(5 )41) Ξ:(ΚD /(∗1/+&%& )( )4% (.)%∃ &.∃,1∗% (, )4% −%+()+∗ &∋+∀2/%
∋(/% =(2: ΒWΧεΓ0 54%∃% +) .∀2%∃3(%& ).∃∀(7%∃0 ∃%3./1)%2 =: )4%
−%+(&+& &)1)%8 ⊥% &4(5 )41) Ξ:(ΚD 41& 1∀ +−∋(∃)1∀) ∃(/% +∀
−1+∀)1+∀+∀3 )4% +∀)%3∃+): (, )4% WΧε 1∀2 )4%∃%=: %∀&.∃%& )41) −%+()+∗
&%3∃%31)+(∀ +& ∗((∃2+∀1)%2 5+)4 &∋(∃% 51// &:∀)4%&+&8
Results
Journal of Cell Science
Fission yeast class V myosins recruit to the site of cell fusion
!∀&∋%∗)+(∀ (, )4% ∗yo567 %6∋∃%&&+(∀ ∋∃(,+/% +∀2+∗1)%& )41) 1/)4(.34
+)& )∃1∀&∗∃+∋)+(∀ 2(%& ∀() 71∃: 2.∃+∀3 )4% 7%3%)1)+7% /+,% ∗:∗/%0 .∋(∀
%∀)∃: +∀)( −%+(&+& 1∀2 &∋(∃./1)+(∀0 )4% %6∋∃%&&+(∀ (, ∗yo567
+∀∗∃%1&%& 2∃1−1)+∗1//: ΒΞ1)1 %) 1/80 ΜΝΝΜϑ Τ.&)+∗+ %) 1/80 ΜΝΝΣΓ8 ⊥%
)4%∃%,(∃% %61−+∀%2 54%)4%∃ )4% &.=&%<.%∀) ∋∃()%+∀ 412 1∀:
2+&∗%∃∀+=/% /(∗1/+&1)+(∀ ∋1))%∃∀ 2.∃+∀3 )4% −%+()+∗ /+,%∗:∗/%8 9(
,1∗+/+)1)% )4+&0 1 &%∃+%& (, &)∃1+∀& %6∋∃%&&+∀3 Ξ:(ΚD≅,/.(∃(∋4(∃%
,.&+(∀& 5%∃% 3%∀%∃1)%28 94% ,/.(∃(∋4(∃% (∃ +)& ∋(&+)+(∀ (∀ Ξ:(ΚD
412 ∀( 2+&∗%∃∀+=/% %,,%∗) .∋(∀ ∗%// −(∃∋4(/(3: 1∀2 %1∗4 ,.&+(∀
∋∃()%+∀ /(∗1/+&%2 )( )4% ∗:)(>+∀%)+∗ 1∗)(−:(&+∀ ∃+∀3 ΒΡΤΓ 2.∃+∀3
−+)(&+& ΒΑ+38 DΓ0 1& ∋∃%7+(.&/: ∃%∋(∃)%2 Β⊥+∀ %) 1/80 ΜΝΝDΓ8 φ+7%≅
∗%// +−13+∀3 (, ∀+)∃(3%∀≅&)1∃7%2 4(−()41//+∗ ∗yo568∋93 4ΕΝ ∗%//&
.∀2%∃3(+∀3 ∋4%∃(−(∀%≅+∀2.∗%2 ∋(/1∃+&1)+(∀ 1∀2 ∗%// ,.&+(∀ ∃%7%1/%2
)41) Ξ:(ΚD )∃1∀&+%∀)/: /(∗1/+&%2 )( )4% /%12+∀3 %23% (, ∋(/1∃+&%2
∗%// )+∋& (, 133/.)+∀1)+∀3 ∗%//&0 +−−%2+1)%/: =%,(∃% ∗%// ,.&+(∀ ΒΑ+38
Dε0 _Κ −+∀.)%&Γ8 94%&% ,(∗+ ∗(1/%&∗%2 +∀)( 1 &+∀3/% =∃+34) ∋(+∀)
ΒΑ+38 Dε0 ΥΚ≅DΜΝ −+∀.)%&Γ0 54+∗4 ∋%∃&+&)%2 .∀)+/ ∗(−∋/%)+(∀ (, ∗%//
51// 2%3∃121)+(∀0 54%∀ )4% Ξ:(ΚD &+3∀1/ 2+&∋%∃&%2 ΒΑ+38 Dε0 DΙΝ≅
DΙΚ −+∀.)%&Γ8
94% 1∗)+∀ 1∀2 −+∗∃().=./% ∗:)(&>%/%)(∀& 417% >%: ∃(/%& +∀ )4%
∗((∃2+∀1)+(∀ 1∀2 %6%∗.)+(∀ (, ∗%// ,.&+(∀ +∀ ,+&&+(∀ :%1&) ΒΧ%)%∃&%∀
%) 1/80 DΕΕΦ1ϑ Χ%)%∃&%∀ %) 1/80 DΕΕΦ=Γ8 ⊥% )4%∃%,(∃% %6∋/(∃%2 )4%
∃%/+1∀∗% (, Ξ:(ΚD .∋(∀ )4% +∀)%3∃+): (, %1∗4 ∗:)(&>%/%)(∀ ,(∃
∃%∗∃.+)−%∀) )( )4% ,.&+∀3 )+∋8 22+)+(∀ (, ∗1∃=%∀21;+− ΒΡεγΓ
∃%&./)%2 +∀ )4% ∃1∋+2 2%∋(/:−%∃+&1)+(∀ (, −+∗∃().=./%&0 =.) 412 ∀(
%,,%∗) .∋(∀ )4% /(∗1/+&1)+(∀ (, Ξ:(ΚD )( )4% &+)% (, ∗%// ,.&+(∀ ΒΑ+38
DΡ0δΓ8 ε: ∗(∀)∃1&)0 2%∋(/:−%∃+&1)+(∀ (, )4% 1∗)+∀ ∗:)(&>%/%)(∀ =:
)4% 122+)+(∀ (, /1)∃.∀∗./+∀  ∀() (∀/: ∋∃%7%∀)%2 ∗%// ,.&+(∀ ,∃(−
)1>+∀3 ∋/1∗%0 =.) 1/&( 1=(/+&4%2 =()4 1∗)+∀ 1∀2 Ξ:(ΚD /(∗1/+&1)+(∀
ΒΑ+38 D]0ΑΓ0 +∀2+∗1)+∀3 )41) )4% ∃%∗∃.+)−%∀) (, )4+& −()(∃ )( )4% &+)%
(, ∗%// ,.&+(∀ ∃%<.+∃%& 1∀ 1∗)+∀ ∗:)(&>%/%)(∀8
⊥% ∀%6) 2%)%∃−+∀%2 54%)4%∃ Ξ:(ΚD ∃%∗∃.+)−%∀) )( )4% ∋∃(?%∗)+(∀
)+∋ ∃%<.+∃%2 Ξ:(ΚΜ0 )4% ()4%∃ ,+&&+(∀ :%1&) ∗/1&& α −:(&+∀ ΒΑ+38
DςΓ8 /)4(.34 ∗yo568∋93 ∗yo5:! ∗%//& 2+&∋/1:%2 )4% 3∃(5)4
∋(/1∃+): 2%,%∗)& 1&&(∗+1)%2 5+)4 )4% 1=&%∀∗% (, Ξ:(ΚΜ0 Ξ:(ΚD 2+2
∀() ∃%<.+∃% Ξ:(ΚΜ ,(∃ +)& /(∗1/+&1)+(∀ 2.∃+∀3 ∗%// ,.&+(∀8 Η(5%7%∃0
1/)4(.34 )4%∃% +& ∀( .∋∃%3./1)+(∀ (, Ξ:(ΚΜ %6∋∃%&&+(∀ 2.∃+∀3
−%+(&+& ΒΞ1)1 %) 1/80 ΜΝΝΜΓ0 )4+& ∋∃()%+∀ 51& 1/&( &%%∀ )( /(∗1/+&%
)( )4% &+)% (, ∗%// ,.&+(∀ ΒΑ+38 DΗΓ +∀ 1∀ 1∗)+∀≅2%∋%∀2%∀) −1∀∀%∃
1∀2 +∀2%∋%∀2%∀)/: (, Ξ:(ΚD ΒΑ+38 D!Γ8 94% /(∗1/+&1)+(∀ ∋1))%∃∀ ,(∃
Ξ:(ΚD 1∀2 Ξ:(ΚΜ 2.∃+∀3 −1)+∀3 51& ∃%−+∀+&∗%∀) (, )41) &%%∀ ,(∃
)4% ,+&&+(∀ :%1&) ,(∃−+∀ Α.&D ΒΧ%)%∃&%∀ %) 1/80 DΕΕΦ∗Γ0 1 ∋∃()%+∀ )41)
+& ∃%<.+∃%2 ,(∃ ∋(/:−%∃+&1)+(∀ (, 1∗)+∀ ,+/1−%∀)& 1) )4% ∋∃(?%∗)+(∀
)+∋ (, −1)+∀3 ∗%//&8 Ο∋(∀ +∀7%&)+31)+(∀0 %1∗4 −:(&+∀ α ∋∃()%+∀ ,1+/%2
Fig. 1. Ξ:(ΚD /(∗1/+&%& )( )4% &+)% (, ∗%// ,.&+(∀8 ΒΓ!Ξ:(ΚD /(∗1/+&%& )( )4% ∗%//
%<.1)(∃ (, −+)()+∗ ∗%//& 54%∀ )133%2 1) %+)4%∃ +)& Ψ≅)%∃−+∀.& ΒςΑΧ≅Ξ:(ΚD 1∀2
Η∗Τ%2D≅Ξ:(ΚDΓ (∃ Ρ≅)%∃−+∀.& ΒΞ:(ΚD≅ςΑΧ 1∀2 Ξ:(ΚD≅Ρ4%∃∃:Γ8 ΒεΓ!9+−%≅
/1∋&% +−13+∀3 (, −%∃3%2 0≅&%∃+%& ∃%7%1/& )41) Ξ:(ΚD )∃1∀&+%∀)/: /(∗1/+&%& )( )4%
&+)% (, ∗%// ,.&+(∀ (, −1)+∀3 ∗%//&8 ΒΡ0δΓ!W+−./)1∀%(.& 7+&.1/+&1)+(∀ (, Ξ:(ΚD
Β−13%∀)1Γ 1∀2 −+∗∃().=./%& Β3∃%%∀Γ +∀ /+7% ∗%//& )∃%1)%2 5+)4 %+)4%∃ δΞW∴ ΒΡΓ
(∃ ΜΚ!∀3χ−/ Ρεγ ΒδΓ &4(5 )41) +)& /(∗1/+&1)+(∀ +& ∀() −+∗∃().=./% 2%∋%∀2%∀)8 !∀
∗(∀)∃1&) )( δΞW∴ )∃%1)−%∀) Β]Γ0 Ξ:(ΚD Β−13%∀)1Γ 1∀2 1∗)+∀ Β3∃%%∀Γ
/(∗1/+&1)+(∀ +& 1=(/+&4%2 +∀ ,.&+∀3 ∗yo568c/erry ∋938&ct6 ∗%//& )∃%1)%2 5+)4
ΜΝ!∀Ξ φ1)  ΒΑΓ8 ΒςΓ!Ξ:(ΚD /(∗1/+&%& )( )4% ,.&+∀3 )+∋ +∀ )4% 1=&%∀∗% (, Ξ:(ΚΜ8
Ξ:(ΚΜ /(∗1/+&%& )( )4% &+)% (, ∗(∀?.31)+(∀ +∀ ∗%//& %+)4%∃ ∋(&&%&&+∀3 ΒΗΓ (∃
/1∗>+∀3 Β!Γ Ξ:(ΚD ΒΞ:(ΚΜ0 3∃%%∀ϑ ∋41&%≅∗(∀)∃1&)0 3∃%:Γ8 ΒηΓ!Ξ:(ΚD ,1+/& )(
/(∗1/+&% )( )4% )+∋ (, 9us6! ∗%//&0 54%∃%1& ∋41&%≅∗(∀)∃1&) +−13%& ∃%7%1/ )41) 1 /1∗>
(, Ξ:(ΚD ΒΛ0 ∗yo56! 9us6!Γ 2(%& ∀() 1//%7+1)% )4% )+∋ ,.&+(∀ ∋4%∀():∋% (, ∗%//&
/1∗>+∀3 )4+& ,(∃−+∀ Βφ0 9us6!Γ8 W∗1/% =1∃β Κ!∀−8
4332
Journal of Cell Science 122 (23)
)( /(∗1/+&% )( )4% ,.&+(∀ &+)% +∀ 133/.)+∀1)+∀3 9us6! ∗%//& ΒΑ+38 DηΓ0
+∀2+∗1)+∀3 )41) )4%: %1∗4 ∃%<.+∃% Α.&D≅1∗)+71)%2 1∗)+∀ ,+/1−%∀)& )(
/(∗1/+&% )( )4% &+)% (, ∗(∀?.31)+(∀8 !∀)%∃%&)+∀3/: )4% %6)%∀2%2
∋∃(?%∗)+(∀ )+∋ ∋4%∀():∋% &%%∀ +∀ 9us6! ∗%//& ΒΧ%)%∃&%∀ %) 1/80 DΕΕΚΓ
ΒΑ+38 DΛΓ 51& &)+// (=&%∃7%2 +∀ 9us6! ∗yo56! ∗%//& ΒΑ+38 DφΓ8
Journal of Cell Science
Overexpression of the GTD of S. pombe myosin V disrupts cell
fusion
Ξ:(&+∀ α ∋∃()%+∀& =+∀2 )4%+∃ ∗1∃3(%& )4∃(.34 )4%+∃ 3/(=./1∃ )1+/
2(−1+∀ Βς9δΓ ΒΤ%∗>≅Χ%)%∃&(∀ %) 1/80 DΕΕΕΓ8 ⊥% )4%∃%,(∃%
3%∀%∃1)%2 &)∃1+∀& +∀ 54+∗4 ςΑΧ 51& ,.&%2 )( )4% Ψ≅)%∃−+∀.& (,
Ξ:(ΚD≅ς9δ 1∀2 %61−+∀%2 )4% %,,%∗) +)& (7%∃%6∋∃%&&+(∀ 412 .∋(∀
∗%// ,.&+(∀8 φ(5≅/%7%/ %6∋∃%&&+(∀ (, Ξ:(ΚD≅ς9δ 412 ∀(
2+&∗%∃∀+=/% %,,%∗) .∋(∀ )4% /(∗1/+&1)+(∀ (, )4% %∀2(3%∀(.& ,.//≅
/%∀3)4 ∋∃()%+∀ +∀ 7%3%)1)+7% ΒΑ+38 ΜΓ (∃ ∗(∀?.31)+∀3 ∗%//& ΒΑ+38
ΜεΓ8 ∴7%∃%6∋∃%&&+(∀ (, )4% Ξ:(ΚD≅ς9δ 412 ∀( 2%)%∗)1=/% %,,%∗)
+∀ 7%3%)1)+7% ∗%//&0 =.) 412 1 2+&∗∃%)% /(∗1/+&1)+(∀ 1& 1 &+∀3/% =∃+34)
2() 1&&(∗+1)%2 5+)4 )4% ∃%3+(∀ (, )4% ∗%// −%−=∃1∀% ∗/(&%&) )( )4%
∀.∗/%.& ΒΑ+38 ΜΡΓ8 ⊥4%∀ (7%∃%6∋∃%&&%2 +∀ ∗(∀?.31)+∀3 ∗%//&0 )4%
Ξ:(ΚD≅ς9δ ,(∃−%2 1 ∃1∀2(−/: /(∗1/+&%2 ∋/1<.%≅/+>% &)∃.∗).∃%0
54+∗4 2+&∋/1∗%2 )4% ,.//≅/%∀3)4 ∋∃()%+∀ ,∃(− )4% &+)% (, ∗%// ,.&+(∀
ΒΑ+38 ΜδΓ0 1∀2 ∋∃%7%∀)%2 ∀(∃−1/ ∗%// ,.&+(∀ =%)5%%∀ )4% )5(
−1)+∀3 ∗%//& ΒΑ+38 Μ]Γ8 /)4(.34 )4+& 2+2 ∀() ∋∃%7%∀) )4%&%
;:3()%& ,∃(− .∀2%∃3(+∀3 ∀.∗/%1∃ ,.&+(∀ 1∀2 −%+(&+&0 )4% /1∗> (,
∗%// 51// %6∋1∀&+(∀ 1) )4% &+)% (, ,.&+(∀ (,)%∀ ∋∃%7%∀)%2 )4%
−(7%−%∀) (, )4% /1∃3% 2+∋/(+2 ∀.∗/%.& =%)5%%∀ )4% )5( ∗%//
∗(−∋1∃)−%∀)&8 94+& ∃%&./)%2 +∀ −%+()+∗ &%3∃%31)+(∀ (,
∗4∃(−(&(−% 1∀2 &∋(∃% ,(∃−1)+(∀0 54+∗4 (∗∗.∃∃%2 +∀ (∀/: (∀% (,
)4% )5( −1)+∀3 ∗%//&0 ∃1)4%∃ )41∀ )4% 54(/% ;:3()% ΒΑ+38 ΜΑ≅!Γ8 ε:
∗(∀)∃1&)0 54%∀ Ξ:(ΚD≅ς9δ 51& (7%∃%6∋∃%&&%2 +∀ ∗(∀?.31)+∀3
∗%//& (∀ ∋/1)%&0 )4%: ,1+/%2 )( ,.&% 1∀2 +∀&)%12 2%7%/(∋%2 %/(∀31)%2
&4−((+∀3 )+∋& ΒΑ+38 ΜηΓ )41) 5%∃% &+−+/1∃ )( )4% &)∃.∗).∃%& (=&%∃7%2
.∋(∀ ,1+/%2 ∗(∀?.31)+(∀ +∀ 9us6! ∗%//& ΒΧ%)%∃&%∀ %) 1/80 DΕΕΚΓ8
Myo51 localises to the meiotic SPB
9( %61−+∀% 54%)4%∃ Ξ:(ΚD 1&&(∗+1)%2 5+)4 1∗)+∀ &)∃.∗).∃%& 2.∃+∀3
−%+()+∗ ∗4∃(−(&(−% &%3∃%31)+(∀0 Ξ:(ΚD≅ςΑΧ /(∗1/+&1)+(∀ 51&
,(//(5%2 +∀ ∗(∀?.31)+∀3 ∗%//& %6∋∃%&&+∀3 1 /1=%//%2 WΧε ∗(−∋(∀%∀)
ΒW+2ΣΓ )( 1//(5 )4% −%+()+∗ &)13% )( =% 2%)%∃−+∀%2 ΒΑ+38 ΙΓ8 & ∀()%2
%1∃/+%∃0 Ξ:(ΚD 51& &%%∀ /(∗1/+&%2 )( )4% &4−(( )+∋ ΒΑ+38 Ι0 ΝβΝΝΓ0
=.) )4+& &+3∀1/ ∃1∋+2/: 2+&∋%∃&%2 .∋(∀ ∗%// ,.&+(∀8 94% ∋∃()%+∀ 412
∀( 2+&∗∃%)% /(∗1/+&1)+(∀ 2.∃+∀3 &.=&%<.%∀) ∀.∗/%1∃ ,.&+(∀ 1∀2
4(∃&%)1+/ −(7%−%∀) ΒΤ(=+∀(50 DΕΥΥΓ Β&.∋∋/%−%∀)1∃: −1)%∃+1/ Α+38
WD0 ΝβΝΚ≅ΣβΙΚΓ8 ∴∀∗% ∗%//& %∀)%∃%2 −%+()+∗ ∋∃(∋41&%0 4(5%7%∃0
Ξ:(ΚD ∃%∗∃.+)%2 )( )4% WΧε0 54%∃% +) ∃%−1+∀%2 /(∗1/+&%2 )4∃(.34(.)
=()4 −%+()+∗ ∗:∗/%& ΒΑ+38 Ι0 ΣβΣΚ≅ΚβΚΚΓ8 Ο∋(∀ %∀)∃: +∀)( −%+(&+& !!0
)4% +∀)%∀&+): (, )4% Ξ:(ΚD &+3∀1/ 1) )4% WΧε +∀∗∃%1&%2 2∃1−1)+∗1//:
ΒΑ+38 Ι0 ΚβΜΚ≅ΚβΙΚΓ0 +∀2+∗1)+∀3 )41) &+3∀+,+∗1∀)/: −(∃% ∋∃()%+∀ 51&
∃%∗∃.+)%2 )( )4% WΧε8 94% &+3∀1/ +∀)%∀&+): (, Ξ:(ΚD 1) )4% WΧε
∃%−1+∀%2 4+34 )4∃(.34(.) ∀.∗/%1∃ &%3∃%31)+(∀ (, −%+(&+& !!0
54%∃%.∋(∀ +) ∃1∋+2/: 2+&1∋∋%1∃%28 δ.∃+∀3 1∀1∋41&% (, −%+(&+& !!0
)4% Ξ:(ΚD &+3∀1/ 51& 1/&( &%%∀ )( 1&&(∗+1)% )( 1 2+&∗∃%)% &)∃.∗).∃%
Fig. 2. Ξ:(&+∀ α ς9δ& ,1+/ )( /(∗1/+&% ∗(∃∃%∗)/: 1∀2 +∀4+=+)
∗%// ,.&+(∀8 Β0εΓ!!∀ ∗(∀)∃1&) )( )4% ,.//≅/%∀3)4 ∋∃()%+∀
Β−13%∀)1Γ0 )4% Ξ:(ΚD≅ς9δ Β3∃%%∀Γ ,1+/& )( /(∗1/+&% +∀ %+)4%∃
−+)()+∗ ΒΓ (∃ ∗(∀?.31)+∀3 ΒεΓ ∗yo568c/erry
n∗t=6∋93∗yo568>?≅ ∗%//& 3∃(5∀ +∀ )4% ∋∃%&%∀∗% (,
)4+1−+∀%8 ΒΡΓ!⊥4%∀ (7%∃%6∋∃%&&%2 +∀ 7%3%)1)+7% ∗%//& Β∀(
)4+1−+∀%Γ0 Ξ:(ΚD≅ς9δ Β−13%∀)1Γ +& ∃%∗∃.+)%2 )( 1 −%2+1/
2() ∀() 1&&(∗+1)%2 5+)4 )4% ∀.∗/%.& Β=/.%Γ8 ΒδΓ!∴7%∃%6∋∃%&&%2
Ξ:(ΚD≅ς9δ Β3∃%%∀Γ /(∗1/+&%& )( 1 ∗:)(∋/1&−+∗ 2() 1∀2
∗1.&%& −+&/(∗1/+&1)+(∀ (, ,.//≅/%∀3)4 Ξ:(ΚD Β−13%∀)1Γ +∀
∗(∀?.31)+∀3 ∗%//&8 Β]0ΑΓ!!∀ )4% 1=&%∀∗% (, 2+&∗∃%)% /(∗1/+&1)+(∀
Β.∋∋%∃ ∋1∀%/&Γ0 ∗%//& (7%∃%6∋∃%&&+∀3 Ξ:(ΚD≅ς9δ ,1+/ )( ,.&%
∀(∃−1//: Β]Γ0 5+)4 &∋(∃%& Β1∃∃(5&Γ ,(∃−+∀3 +∀ (∀/: (∀% (, )4%
)5( −1)+∀3 ∗%//& ΒΑΓ8 ΒςΓ!94% ∋∃(∋(∃)+(∀ (,
%eu6::n∗t=6∗yo568>?≅ ∗%//& 5+)4 ∀(∃−1/ 1∀2 1=∀(∃−1/ 1&∗+
54%∀ 1//(5%2 )( ∗(∀?.31)% +∀ )4% ∋∃%&%∀∗% Β3∃%: =1∃&Γ (∃
1=&%∀∗% Β=/1∗> =1∃&Γ (, Σ!∀Ξ )4+1−+∀%8 ΒΗ0!Γ!W+2Σ≅)(−1)(
/1=%//%2 WΧε& Β−13%∀)1Γ ∃%7%1/ )41) 1/)4(.34 ∀.∗/%1∃ ,.&+(∀
+& .∀1,,%∗)%2 +∀ ΕΝι (, ∗%//& (7%∃%6∋∃%&&+∀3 )4% Ξ:(ΚD≅
ς9δ Β3∃%%∀Γ0 ∀.∗/%1∃ 2+&)∃+=.)+(∀ +& 2+&∃.∋)%2 +∀ )4%
&.=&%<.%∀) −%+(&+&8 ΒηΓ!∴7%∃%6∋∃%&&+(∀ (, Ξ:(ΚD≅ς9δ ∗1∀
=∃+∀3 1=(.) )4% 9us6! ∋4%∀():∋% +∀ ∗%//& (∀ ∋/1)%&8 W∗1/%
=1∃&β Κ!∀−8
Myosin V and fission yeast meiosis
=%4+∀2 )4% ∀.∗/%.& ΒΑ+38 Ι 1∃∃(54%12&Γ0 ∃%−+∀+&∗%∀) (, )4% 1∗)+∀
/(∗1/+&1)+(∀ 1) )4+& )+−% ΒΧ%)%∃&%∀ %) 1/80 DΕΕΦ=Γ8
9( ∗(∀,+∃− )4% WΧε 1&&(∗+1)+(∀ (, Ξ:(ΚD0 +)& /(∗1/+&1)+(∀ 51&
%61−+∀%2 +∀ 1 &)∃1+∀ %6∋∃%&&+∀3 Η∗Τ%2D≅Ξ:(ΚD 1∀2 Χ∗∋D≅ςΑΧ0
54+∗4 +& 1∀ %&&%∀)+1/ WΧε ∗(−∋(∀%∀) ΒΑ/(∃: %) 1/80 ΜΝΝΜΓ8 Ρ(∀&+&)%∀)
5+)4 )4% Ρ≅)%∃−+∀1/≅)133%2 ∋∃()%+∀0 Η∗Τ%2D≅Ξ:(ΚD /(∗1/+&%2 )( )4%
−%+()+∗ WΧε ,∃(− )4% (∀&%) (, ∗4∃(−(&(−% &%3∃%31)+(∀ (, −%+(&+&
! ΒΑ+38 ΣΓ8 94% &+3∀1/& ∗(∀)+∀.%2 )( ∗(1/%&∗% )4∃(.34(.) −%+(&+&
! ΒΑ+38 ΣεΓ0 .∀)+/ )4% %∀2 (, )%/(∋41&% (, −%+(&+& !! ΒΑ+38 ΣΡΓ0 54%∀
)4% &+3∀1/ ,∃(− )4% )5( ∋∃()%+∀& 2+2 ∀() (7%∃/1∋ ∋∃%∗+&%/:0 +∀2+∗1)+∀3
)41) Ξ:(ΚD +& /(∗1)%2 (∀ )4% ∋%∃+∋4%∃: (, )4% WΧε8 Ρ(∀&+&)%∀) 5+)4
)4+&0 1 31∋ 51& (,)%∀ (=&%∃7%2 =%)5%%∀ )4% Ξ:(ΚD &+3∀1/ 1∀2 )4%
3∃(5+∀3 −%+()+∗ &∋+∀2/% ΒΑ+38 ΣδΓ0 )4% &+;% (, 54+∗4 51& ∗(−∋1∃1=/%
5+)4 )4% 2+&)1∀∗% =%)5%%∀ )4% Ξ:(ΚD &+3∀1/ 1∀2 )41) (, )4%
>+∀%)(∗4(∃% ∗(−∋(∀%∀)0 Ξ1/Μ ΒΑ+38 Σ]Γ Βη+∀ %) 1/80 ΜΝΝΜΓ0 1∀2
+∀2+∗1)% )41) Ξ:(ΚD +& /(∗1)%2 )( )4% (.)%∃ ∃%3+(∀ (, )4% WΧε
)4∃(.34(.) −%+()+∗ ∗4∃(−(&(−% &%3∃%31)+(∀8 94+& 2+&)∃+=.)+(∀ 51&
∗(∀,+∃−%2 =: %61−+∀+∀3 )4% Ξ:(ΚD &+3∀1/ +∀ −(∃% )41∀ DΝΝ ∗%//&
,∃(− %1∗4 &)13% (, )4% −%+()+∗ ∗%// ∗:∗/%8 & )4% 2+&)∃+=.)+(∀ 51&
∗(∀&+&)%∀) ,(∃ 1// )4% ∗%//& 5% (=&%∃7%20 5% ∗(∀∗/.2%2 )41) +) ∃%,/%∗)%2
)4% ∀(∃−1/ ∗%//./1∃ 2+&)∃+=.)+(∀ ,(∃ Ξ:(ΚD 2.∃+∀3 −%+(&+&8
Journal of Cell Science
Myo52 concentrates near the SPB and nuclei during meiosis
⊥% ∀%6) %61−+∀%2 54%)4%∃ )4% −1?(∃ −+)()+∗ ∗:∗/% −:(&+∀ α ∋∃()%+∀
Ξ:(ΚΜ0 ∗(/(∗1/+&%2 5+)4 Ξ:(ΚD )( )4% −%+()+∗ WΧε8 δ.∃+∀3 )4%
,+∃&) −%+()+∗ 2+7+&+(∀0 )4% Ξ:(ΚΜ &+3∀1/ ∗(∀∗%∀)∃1)%2 )( 1 ∀.−=%∃
(, 2+&∗∃%)% ,(∗+ ΒΑ+38 ΣΑ0ςΓ &(−% (, 54+∗4 5%∃% 12?1∗%∀) )(0 =.)
∀%7%∃ ∗(/(∗1/+&%2 5+)40 Ξ:(ΚD 1) )4% WΧε ΒΑ+38 Ση0ΛΓ8 δ.∃+∀3 )4+&
∋%∃+(20 ()4%∃ Ξ:(ΚΜ ,(∗+ 1∋∋%1∃%2 )( =% ∃1∀2(−/: 2+&)∃+=.)%2
)4∃(.34(.) )4% ∗:)(∋/1&−8 Ο∋(∀ ∗(−∋/%)+(∀ (, −%+(&+& !0 Ξ:(ΚΜ
/(∗1/+&%2 1∃(.∀2 )4% ∀.∗/%1∃ ∋%∃+∋4%∃: ΒΑ+38 ΣΗΓ0 +∀ 1 ∋1))%∃∀ )41)
51& &+−+/1∃ )( )41) &%%∀ ,(∃ 1∗)+∀ 1) )4+& &)13% ΒΧ%)%∃&%∀ %) 1/80 DΕΕΦ=Γ8
4333
94+& Ξ:(ΚΜ 2+&)∃+=.)+(∀ ∃1∋+2/: 2+&∋%∃&%2 1,)%∃ ∗4∃(−(&(−%
&%3∃%31)+(∀0 1∀2 Ξ:(ΚΜ 51& &.=&%<.%∀)/: (∀/: &%%∀ /(∗1/+&%2 )(
∗:)(∋/1&−+∗ ,(∗+ ΒΑ+38 Σ!Γ8 94%∃%,(∃%0 .∀/+>% Ξ:(ΚD0 2.∃+∀3 −%+(&+&
Ξ:(ΚΜ 2+2 ∀() /(∗1/+&% )( )4% WΧε0 =.) 412 1 2+&)∃+=.)+(∀ &+−+/1∃
)( )41) (, 1∗)+∀8
Myo51 associates with the SPB independently of Myo52,
Spo15, actin and the SIN network
⊥% ∀%6) %61−+∀%2 54%)4%∃ Ξ:(ΚD 51& 2%∋%∀2%∀) .∋(∀ ()4%∃
∗%//./1∃ ∗(−∋(∀%∀)& ,(∃ +)& /(∗1/+&1)+(∀ )( )4% −%+()+∗ WΧε8 ⊥% ,+∃&)
%&)1=/+&4%2 )41) Ξ:(ΚD ∃%∗∃.+)%2 )( )4% WΧε +∀ )4% 1=&%∀∗% (, )4%
&%∗(∀2 ,+&&+(∀ :%1&) ∗/1&& α −:(&+∀ Ξ:(ΚΜ ΒΑ+38 ΣφΓ8 W+−+/1∃/:0
W∋(DΚ0 54+∗4 +& ∃%<.+∃%2 ,(∃ −(2+,+∗1)+(∀ (, )4% WΧε 2.∃+∀3 &∋(∃%
−%−=∃1∀% ,(∃−1)+(∀ Β!>%−()( %) 1/80 ΜΝΝΝΓ 51& ,(.∀2 )( =%
2+&∋%∀&1=/% ,(∃ ∃%∗∃.+)−%∀) 1∀2 /(∗1/+&1)+(∀ (, Ξ:(ΚD )( )4% WΧε
ΒΑ+38 ΣΞΓ8 Ρ(∃% ∗(−∋(∀%∀)& (, )4% ,+&&+(∀ :%1&) &%∋)1)+(∀ +∀+)+1)+(∀
∀%)5(∃> ΒW!ΨΓ 1∃% 1∀∗4(∃%2 )( )4% (.)%∃ ∋/1<.% (, )4% WΧε =: Ρ2∗DD
ΒΛ∃1∋∋ %) 1/80 ΜΝΝDΓ0 )( ∃%3./1)% &∋(∃% ,(∃−1)+(∀ 2.∃+∀3 −%+(&+&
ΒΛ∃1∋∋ %) 1/80 ΜΝΝ_Γ8 Ο&+∀3 1 )%−∋%∃1).∃%≅&%∀&+)+7% c,c66 −.)1∀)0
5% 1&&%&&%2 54%)4%∃ Ξ:(ΚD 51& 1∀∗4(∃%2 )( )4% WΧε 7+1 Ρ2∗DD0
(∃ 54%)4%∃ W!Ψ 1∗)+7+): +& ∃%<.+∃%2 ,(∃ +)& /(∗1/+&1)+(∀8 ⊥4%∀ c,c668
6Β: ∗yo568∋93 ∗%//& 5%∃% ∗./).∃%2 1) ΙΙϕΡ Β∀(∀≅∋%∃−+&&+7%
)%−∋%∃1).∃%Γ0 Ξ:(ΚD 51& &%%∀ )( /(∗1/+&% )( )4% ∀(∀≅∗(∀&)∃+∗)+∀3
ΡΤ 2.∃+∀3 −+)(&+&0 1∀2 1/&( )( )4% WΧε& )4∃(.34(.) −%+(&+& ΒΑ+38
ΣΨΓ0 +∀2+∗1)+∀3 )41) Ξ:(ΚD /(∗1/+&1)+(∀ +& +∀2%∋%∀2%∀) (, Ρ2∗DD
(∃ W!Ψ 1∗)+7+):8
9( %6∋/(∃% )4% 2%∋%∀2%∀∗% (, Ξ:(ΚD .∋(∀ 1∗)+∀ ,(∃ +)&
/(∗1/+&1)+(∀ 2.∃+∀3 −%+(&+&0 5% ∗∃%1)%2 1 &)∃1+∀ ∗(∀)1+∀+∀3 )4%
∗yo568∗Χ/erry 1//%/% )(3%)4%∃ 5+)4 1 ∋938&ct6 1//%/% .∀2%∃
)4% ∗(∀)∃(/ (, )4% n∗tD6 ∋∃(−()%∃0 54+∗4 412 =%%∀ +∀)%3∃1)%2 +∀)(
)4% %eu67 /(∗.&8 −Ρ4%∃∃:≅/1=%//%2 Ξ:(ΚD /(∗1/+&%2 )( )4% WΧε& +∀
)4% ∋∃%&%∀∗% (, δΞW∴ ΒΑ+38 Σ∴Γ8 Η(5%7%∃0 1/)4(.34 ,/.(∃%&∗%∀∗%
,∃(− ςΑΧ≅1∗)+∀ ,(∗+ 51& ∃1∋+2/: ∃%∋/1∗%2 5+)4 1 ∀(∀≅2+&∗∃%)%
∗:)(∋/1&−+∗ =1∗>3∃(.∀2 &+3∀1/ .∋(∀ 122+)+(∀ (, )4% 1∗)+∀≅
2%∋(/:−%∃+&+∀3 2∃.3 /1)∃.∀∗./+∀  ΒΑ+38 ΣΧΓ0 Ξ:(ΚD ∃%−1+∀%2
/(∗1/+&%2 1) )4% WΧε0 &.33%&)+∀3 )41) )4+& /(∗1/+&1)+(∀ 51&
+∀2%∋%∀2%∀) (, +)& 1&&(∗+1)+(∀ 5+)4 1∗)+∀ 1∀2 )4%∃%,(∃% −()(∃
1∗)+7+):8
Myo51 turns over on the SPB during meiosis I
Η17+∀3 %&)1=/+&4%2 )41) Ξ:(ΚD 2(%& ∀() ∃%<.+∃% 1∗)+∀ )(
/(∗1/+&% )( )4% WΧε0 5% ∀%6) %61−+∀%2 54%)4%∃ )4+& /(∗1/+&1)+(∀
51& 2:∀1−+∗8 Ο&+∀3 ,/.(∃%&∗%∀∗% ∃%∗(7%∃: 1,)%∃
∋4()(=/%1∗4+∀3 ΒΑΤΧΓ0 5% %61−+∀%2 Ξ:(ΚD≅ςΑΧ ).∃∀(7%∃
1) )4% WΧε (, ∗%//& +∀ −%+(&+& ! ΒΑ+38 ΚΓ (∃ −%+(&+& !! ΒΑ+38
ΚΡ0]Γ8 δ.∃+∀3 −%+(&+& !0 )4%∃% 51& 1 ∃1∋+2 ∃%∗(7%∃: (, )4%
ςΑΧ≅Ξ:(ΚD &+3∀1/ 1) )4% WΧε 1,)%∃ ∋4()(=/%1∗4+∀30 1∀2 ,.//
∃%∗(7%∃: )((> /%&& )41∀ Κ −+∀.)%& ΒΑ+38 Κ0εΓ8 94%&% ∗%//&
5%∃% )4%∀ 1=/% )( ∗(−∋/%)% −%+(&+& 1∀2 &∋(∃% ,(∃−1)+(∀
∀(∃−1//: Β∀() &4(5∀Γ0 +∀2+∗1)+∀3 )41) )4% /1&%∃ 412 ∀(
2%)%∗)1=/% 1,,%∗) .∋(∀ )4% +∀)%3∃+): (, )4% WΧε8 94% ∃%∗(7%∃%2
Fig. 3. Ξ:(ΚD /(∗1/+&%& )( )4% WΧε 2.∃+∀3 −%+()+∗ ∀.∗/%1∃ &%3∃%31)+(∀8
Ξ16+−.− ∋∃(?%∗)+(∀& (, ΜD &/+∗% 0≅&)1∗>& &%/%∗)%2 ,∃(− 1 )+−% ∗(.∃&% (,
−1)+∀3 ∗yo568∋93 si,=8to∗&to ∗%//& ΒΑ.// )+−% ∗(.∃&% +& &4(5∀ +∀
&.∋∋/%−%∀)1∃: −1)%∃+1/ Α+38 WDΓ ∃%7%1/ )41) Ξ:(ΚD Β3∃%%∀Γ ∀() (∀/:
/(∗1/+&%& )( ∗(∀?.31)+∀3 )+∋& =.) 1/&( ∗(≅/(∗1/+&%& 5+)4 )4% WΧε
∗(−∋(∀%∀) W+2Σ Β−13%∀)1Γ 2.∃+∀3 −%+()+∗ ∗4∃(−(&(−% &%3∃%31)+(∀
ΒΣβΙΚ≅_βΝΝΓ8 ) )4% %∀2 (, −%+(&+& !!0 Ξ:(ΚD )∃1∀&+%∀)/: /(∗1/+&%& )( 1
2+&∋%∃&%2 &)∃.∗).∃% =%4+∀2 %1∗4 WΧε Β1∃∃(54%12& +∀ ΚβΚΝ≅_βΝΝΓ8 9+−% +&
&4(5∀ +∀ 4(.∃&β−+∀.)%&8
Journal of Cell Science 122 (23)
Journal of Cell Science
4334
Fig. 4. Ξ:(ΚD /(∗1/+&%& )( )4% (.)&+2% (, −%+()+∗ WΧε&8 Β≅ΡΓ!Ξ:(ΚD Β−13%∀)1Γ ∗(/(∗1/+&%& 5+)4 )4% %&&%∀)+1/ WΧε ∗(−∋(∀%∀) Χ∗∋D Β3∃%%∀Γ ,∃(− )4% (∀&%) (,
−%+()+∗ ∀.∗/%1∃ &%3∃%31)+(∀ ΒΓ0 )4∃(.34(.) −%+(&+& ! ΒεΓ0 .∀)+/ )4% %∀2 (, −%+(&+& !! ΒΡΓ +∀ ΕcRe,68∗yo56 3c368∋93 ∗%//&8 ΒδΓ! 31∋ +& &%%∀ =%)5%%∀ Ξ:(ΚD
Β−13%∀)1Γ 1∀2 )4% %∀2 (, %/(∀31)+∀3 ςΑΧ≅)=Μ /1=%//%2 −%+()+∗ &∋+∀2/%& Β3∃%%∀Γ Β/(5%∃ +−13%& )1>%∀ Σ −+∀.)%& 1,)%∃ .∋∋%∃ +−13%&Γ &.33%&)+∀3 Ξ:(ΚD /(∗1/+&%& )(
)4% (.)%∃ &+2% (, )4% WΧε8 WΧε ∃%3+(∀& 1∃% −13∀+,+%2 +∀ ∗(−∋(&+)% +−13%&8 Β]Γ!Ξ:(ΚD Β−13%∀)1Γ ,1+/%2 )( ∗(/(∗1/+&% 5+)4 )4% >+∀%)(∗4(∃% ∗(−∋(∀%∀)0 Ξ1/Μ Β3∃%%∀Γ
+∀ ΕcRe,68∗yo56 ∗&%:8∋93 ∗%//&8 ΒΑ≅ΛΓ!Ξ:(ΚD Β3∃%%∀Γ 1∀2 Ξ:(ΚΜ Β−13%∀)1Γ /(∗1/+&1)+(∀ +∀ ∗yo568∋93 ∗yo5:t,?o∗&to ∗%//& 2.∃+∀3 −%+(&+&8 Ξ:(ΚΜ &+3∀1/
∗(∀∗%∀)∃1)%& 12?1∗%∀) )( Ξ:(ΚD≅/1=%//%2 WΧε& 2.∃+∀3 −%+(&+& ! ΒΑ0ςΓ0 1∃(.∀2 )4% ∀.∗/%.& 1) )4% (∀&%) (, −%+(&+& !! ΒΗΓ0 1∀2 &.=&%<.%∀)/: )( ∋.∀∗)1)% ,(∗+ )41) 1∃%
∃1∀2(−/: 2+&)∃+=.)%2 Β!Γ8 Βη0ΛΓ!Ξ13∀+,+%2 WΧε ∃%3+(∀& 4+34/+34)%2 +∀ Α 1∀2 ς0 ∃%&∋%∗)+7%/:8 ΒφΓ!Ξ:(ΚD /(∗1/+&%& )( )4% WΧε (, −%+()+∗ ∗yo5:! ∗%//&8 ΒΞΓ!ςΑΧ
Β.∋∋%∃ ∋1∀%/Γ 1∀2 −%∃3%2 ςΑΧ ∋41&%≅∗(∀)∃1&) +−13% Β/(5%∃ ∋1∀%/Γ ∃%7%1/ )41) Ξ:(ΚD /(∗1/+&%& )( )4% WΧε (, ∋938∗yo6 s3o65! ∗%//& 2.∃+∀3 −%+(&+&8 ΒΨΓ!Ξ:(ΚD
/(∗1/+&1)+(∀ )( )4% ΡΤ Β1∃∃(5Γ (∃ −%+()+∗ WΧε Β1∃∃(54%12&Γ +& ∀(∃−1/ +∀ c,c6686Β: ∗%//& 3∃(5∀ 1) )4% ∀(∀≅∋%∃−+&&+7% )%−∋%∃1).∃%8 Β∴Γ!δΞW∴ 41& ∀( %,,%∗) (∀
Ξ:(ΚD Β−13%∀)1Γ (∃ 1∗)+∀ Β3∃%%∀Γ /(∗1/+&1)+(∀ 2.∃+∀3 −%+(&+&8 ΒΧΓ!9∃%1)−%∀) 5+)4 DΝ!∀Ξ /1)∃.∀∗./+∀  =∃+∀3& 1=(.) ∃1∋+2 2%∋(/:−%∃+&1)+(∀ (, 1∗)+∀0 =.) 41& ∀(
%,,%∗) (∀ Ξ:(ΚD /(∗1/+&1)+(∀ )( )4% WΧε Β1∃∃(54%12&Γ8 W∗1/% =1∃β Κ!∀−8
Myosin V and fission yeast meiosis
ςΑΧ≅Ξ:(ΚD &+3∀1/ 51& ∗(−∋1∃1=/% 5+)4 )41) &%%∀ (∀ )4% &%∗(∀2
.∀=/%1∗4%2 WΧε 1) )4% ()4%∃ %∀2 (, )4% ∗%//0 +∀2+∗1)+∀3 )41) WΧε≅
1&&(∗+1)%2 Ξ:(ΚD +& 7%∃: 2:∀1−+∗ 2.∃+∀3 −%+(&+& !8 ε: ∗(∀)∃1&)0
54%∀ )4% ςΑΧ≅Ξ:(ΚD &+3∀1/ 51& =/%1∗4%2 ,∃(− 1 &+∀3/% WΧε (,
1 ∗%// +∀ −%+(&+& !! ΒΑ+38 ΚΡ0]Γ0 )4% ςΑΧ &+3∀1/ ∀%7%∃ ∃%∗(7%∃%2 )(
∀(∃−1/ /%7%/& ΒΑ+38 ΚδΓ0 %7%∀ 1,)%∃ %6)%∀2%2 )+−%≅/1∋&% 1∀1/:&+&
ΒΑ+38 ΚΑΓ0 =: 54+∗4 )+−% )4% ςΑΧ &+3∀1/ 1) )4% .∀1,,%∗)%2 WΧε& 412
&)1∃)%2 )( 2+&1∋∋%1∃8 94%&% 21)1 +∀2+∗1)% )41) Ξ:(ΚD 41& 2+&)+∀∗)
−%+()+∗≅∗:∗/%≅2%∋%∀2%∀) 2:∀1−+∗ ∋∃(∋%∃)+%& 1) )4% WΧε8
Journal of Cell Science
MPF is required for Myo51 meiosis II SPB recruitment
9( %6∋/(∃% )4% ∋∃%∗+&% )+−+∀3 (, )4% +∀∗∃%1&% +∀ Ξ:(ΚD &+3∀1/ 1)
)4% WΧε 1∀2 1//(5 .& )( ∗(∃∃%/1)% ∗41∀3%& +∀ +)& ).∃∀(7%∃0 5%
%61−+∀%2 Ξ:(ΚD /(∗1/+&1)+(∀ +∀ 1 &)∃1+∀ /1∗>+∀3 Ξ%&D0 54+∗4 +& 1
∃%3./1)(∃ (, −%+(&+& !! (∀&%)8 Ρ%//& /1∗>+∀3 Ξ%&D 1∃∃%&) 1) )4% %∀2
(, −%+(&+& ! =%∗1.&% (, )4% ∋∃%−1).∃% 2%3∃121)+(∀ (, )4% ∋((/ (,
∗:∗/+∀ ε ∃%<.+∃%2 ,(∃ (∀&%) (, −%+(&+& !! Β!;151 %) 1/80 ΜΝΝΚϑ W4+−(21
%) 1/80 DΕΦΚΓ8 & ∋∃%2+∗)%20 ∗es67 ∗%//& .∀2%∃5%∀) ∀(∃−1/ −%+()+∗
WΧε &%3∃%31)+(∀0 5+)4 )4% %6∋%∗)%2 +∀∗∃%1&% +∀ Ξ:(ΚD &+3∀1/ 2.∃+∀3
−%+(&+& !! ΒΑ+38 _Γ8 ε: ∗(∀)∃1&)0 1 ∃1∋+2 ∃%2.∗)+(∀ +∀ Ξ:(ΚD WΧε
&+3∀1/ 51& (=&%∃7%2 +∀ ∗es6! ∗%//& .∋(∀ )4%+∃ 1∃∃%&) 1) )4% %∀2 (,
−%+(&+& ! ΒΑ+38 _εΓ0 1∀2 )4% −()(∃ 51& &%%∀ )( 1&&(∗+1)% 5+)4 )4%
∀.∗/%1∃ −%−=∃1∀%&8 94+& /(∗1/+&1)+(∀ ∋1))%∃∀ 51& (=&%∃7%2 +∀ 1//
∗yo567 ∗%//& %61−+∀%2 )41) 5%∃% ∋1&&+∀3 ,∃(− ∋∃(∋41&% )( )4% (∀&%)
(, 1∀1∋41&% (, −%+(&+& !! ΒΑ+38 _ΡΓ8 94%&% 21)1 &.33%&) )41) )4%
+∀∗∃%1&% +∀ )4% Ξ:(ΚD WΧε &+3∀1/ ∗(∃∃%/1)%& 5+)4 )4% (∀&%) (,
−%+(&+& !!8
9( ∗(∀,+∃− )4+&0 5% 3%∀%∃1)%2 1 ∗es6! &)∃1+∀ +∀ 54+∗4 +) 51&
∋(&&+=/% )( ∃%3./1)% )4% +∀2.∗)+(∀ (, ∀(∀≅2%3∃121=/% ∗:∗/+∀ ε
ΒΡ2∗DΙ≅2%&ΜΓ .&+∀3 1∀ %&)∃12+(/≅∃%3./1)1=/% 4(∃−(∀%≅=+∀2+∀3
2(−1+∀ ΒΗεδΓ Βε(% %) 1/80 ΜΝΝΦΓ0 1∀2 )4%∃%=: +∀∗∃%1&% Ρ2∗DΙ /%7%/&
4335
1) )4% ∗es6!≅+∀2.∗%2 1∃∃%&) ∋(+∀)8 !∀ )4% 1=&%∀∗% (, %&)∃12+(/0 )4%
Ηεδ≅)133%2 ∀(∀≅2%3∃121=/% Ρ2∗DΙ ∋∃()%+∀ +& )1∃3%)%2 )( )4%
Η&∋ΕΝ ∗(−∋/%60 ∃%∀2%∃+∀3 +) +∀1∗)+7%8 Η(5%7%∃0 )4% 122+)+(∀ (,
%&)∃12+(/ =∃+∀3& 1=(.) 1∀ 1/−(&) +∀&)1∀)1∀%(.& ∃%/%1&% (, )4% Ηεδ≅
Ρ2∗DΙ≅2%&Μ ,.&+(∀ ∋∃()%+∀ Βε(% %) 1/80 ΜΝΝΦΓ8 /)4(.34 5% 5%∃%
.∀1=/% )( 2%)%∗) Ξ:(ΚD 1) )4% WΧε (, ∗yo568∋93 ∗es6! ∋Τ]ΧΦΜ
c,c6Β,es:8ΕΓ≅0 ∗(∀)∃(/ ∗%//& )∃%1)%2 5+)4 %)41∀(/ ΒΑ+38 _δ0
1∃∃(54%12&Γ0 .∋(∀ 122+)+(∀ (, %&)∃12+(/0 Ξ:(ΚD 51& &%%∀ )( /(∗1/+&%
)( )4% WΧε ΒΑ+38 _δ =())(− ∋1∀%/&Γ8 94+& ,+∀2+∀3 +& ∗(∀&+&)%∀) 5+)4
)4% +∀∗∃%1&% +∀ Ξ:(ΚD &+3∀1/ 1) )4% WΧε 1∀2 )4% ∗(∃∃%&∋(∀2+∀3
∃%2.∗)+(∀ (=&%∃7%2 +∀ +)& ).∃∀(7%∃ 1,)%∃ )4% (∀&%) (, −%+(&+& !!0
+∀2+∗1)+∀3 )41) Ξ:(ΚD −%+(&+& !! WΧε ∃%∗∃.+)−%∀) ∃%<.+∃%& )4%
∋∃%&%∀∗% (, 1∗)+7% ΞΧΑ8
Modulating Myo51 protein levels disrupts meiotic chromosome
segregation and spore formation
& Ξ:(ΚD 41& 1 2+&)+∀∗) /(∗1/+&1)+(∀ ∋1))%∃∀ 1) )4% WΧε 2.∃+∀3
−%+(&+&0 5% %61−+∀%2 54%)4%∃ Ξ:(ΚD 412 1 ∃(/% 2.∃+∀3 )4% /1))%∃
&)13%& (, −%+(&+& 1∀2 &.=&%<.%∀) &∋(∃% ,(∃−1)+(∀8 Α+∃&)0 5%
%61−+∀%2 )4% ∀.∗/%1∃ &%3∃%31)+(∀ 5+)4+∀ &∋(∃%& ,∃(− 1 4(−(;:3(.&
∗yo56! ∗%// ∗∃(&&8 /)4(.34 %1∗4 &∋(∃% 5+)4+∀ 1&∗+ ,∃(− 1
4(−(;:3(.& ∗∃(&& =%)5%%∀ ∗yo567 ∗%//& ∗(∀)1+∀%2 ∀.∗/%1∃ −1)%∃+1/
ΒΑ+38 ΥΓ0 1 &+3∀+,+∗1∀) ∋∃(∋(∃)+(∀ ΒκDΝιΓ (, 1&∗+ ,∃(− 1 &+−+/1∃
∗∃(&& =%)5%%∀ 4(−(;:3(.& ∗yo56! ∗%//& ∗(∀)1+∀%2 1 &∋(∃% /1∗>+∀3
δΨ ΒΑ+38 Υεϑ 91=/% DΓ0 +∀2+∗1)+7% (, 1 2%,%∗) 2.∃+∀3 −%+(&+& !!8
94+& ,+∀2+∀3 51& ∗(∀,+∃−%2 .&+∀3 1∀ 1//%/% %6∋∃%&&+∀3 1 ςΑΧ ,.&+(∀
(, )4% ∀.∗/%1∃ −%−=∃1∀% ∗(−∋(∀%∀)0 δ/∗Μ ΒΞ+>+ %) 1/80 ΜΝΝΜΓ 1∀2
+//.&)∃1)%2 )41) ∀.∗/%+ (,)%∀ ,1+/%2 )( &%3∃%31)% ∗(∃∃%∗)/: +∀ )4%
1=&%∀∗% (, Ξ:(ΚD ΒΑ+38 ΥΡΓ8 !∀)∃(2.∗+∀3 )4% si,=8t,?o∗&to 1//%/%
+∀)( )4+& &)∃1+∀ ∃%7%1/%2 )41) −%+()+∗ ∗yo56! ∗%//& 412 1∀ +∀∗∃%1&%2
∀.−=%∃ (, WΧε& ΒΑ+38 Υδ0]Γ0 54+∗4 5%∃% (,)%∀ &%%∀ )( 1&&(∗+1)%
5+)4 ,∃13−%∀)& (, ∀.∗/%1∃ −%−=∃1∀% ΒΑ+38 Υ]0
1∃∃(54%12&Γ8 !∀ 122+)+(∀0 1 ∋∃(∋(∃)+(∀ (, )4%
∃%&./)1∀) 1&∗+ 1/&( ∗(∀)1+∀%2 1∀ +∀∗∃%1&%2 ∀.−=%∃
(, &∋(∃%&0 −1∀: (, 54+∗4 ∗(∀)1+∀%2 ∀( (∃ ∃%2.∗%2
<.1∀)+)+%& (, ∀.∗/%1∃ −1)%∃+1/0 54+∗4 51& ∃%,/%∗)%2
+∀ 1 ∃%2.∗)+(∀ +∀ &∋(∃% 7+1=+/+): Β91=/% DΓ8 94%&%
21)1 +∀2+∗1)% )41) Ξ:(ΚD 41& 1 ∃(/% +∀ −1+∀)1+∀+∀3
WΧε +∀)%3∃+): 2.∃+∀3 −%+(&+&8
94+& WΧε +∀)%3∃+): 1∀2 &∋(∃% ,(∃−1)+(∀
∋4%∀():∋%& 5%∃% −(∃% ∋∃(−+∀%∀) +∀ 1&∗+ ∋∃(2.∗%2
,∃(− −1)+∀3 ∗%//& (7%∃%6∋∃%&&+∀3 Ξ:(ΚD ΒΑ+38
ΥΑ0ςϑ 91=/% DΓ8 ∴7%∃%6∋∃%&&+(∀ (, Ξ:(ΚD (,)%∀
∃%&./)%2 +∀ )4% 3%∀%∃1)+(∀ (, 1&∗+ ∗(∀)1+∀+∀3 1∀
1=∀(∃−1/ ∀.−=%∃ (, &∋(∃%& ΒΑ+38 ΥΑ0ςΓ0 54+∗4
&(−%)+−%& ∃%1∗4%2 Ι_8 Η(5%7%∃ WΧε 1∀2 ∀.∗/%1∃
−%−=∃1∀% /(∗1/+&1)+(∀ ∃%7%1/%2 )41) %1∗4 &∋(∃% 2+2
∀() 1/51:& ∗(∀)1+∀ δΨ0 =.) +∀&)%12 ∗(∀)1+∀%2 1∀
1=%∃∃1∀) 1∗∗.−./1)+(∀ (, WΧε −1)%∃+1/ ΒΑ+38 ΥΑΓ0
54+∗4 51& ∃%,/%∗)%2 +∀ )4% 2∃1−1)+∗ ∃%2.∗)+(∀ +∀
Fig. 5. 94% 1−(.∀) (, Ξ:(ΚD 1) )4% −%+()+∗ WΧε +&
2:∀1−+∗8 Β≅ΑΓ!9+−%≅/1∋&% +−13%& 1∀2 3∃1∋4& (, &+3∀1/
∗41∀3% (, Ξ:(ΚD≅ςΑΧ ΑΤΧ %6∋%∃+−%∀)&8 Ξ:(ΚD (∀ )4%
WΧε (, ∗%//& +∀ −%+(&+& ! ΒΓ 1∀2 −%+(&+& !! ΒΡ0]Γ 51&
&.=?%∗)%2 )( ∋4()(=/%1∗4+∀3 Β1∃∃(5&Γ0 1∀2 +)& ∃%∗(7%∃: 51&
,(//(5%2 .&+∀3 Ιδ )+−%≅/1∋&% +−13+∀3 Β/%,) ∋1∀%/&Γ8
ς∃1∋4& Β∃+34)Γ &4(5 ∗41∀3% +∀ ∃%/1)+7% +∀)%∀&+): (, Ξ:(ΚD
&+3∀1/ 1) )4% ∋4()(=/%1∗4%2 Β3∃%: ,+//%2 ∗+∃∗/%&Γ 1∀2 1∀
.∀=/%1∗4%2 WΧε Β%−∋): ∗+∃∗/%&Γ )4∃(.34(.) )4%
%6∋%∃+−%∀)8 9+−% +& ∃%∗(∃2%2 1& %+)4%∃ &%∗(∀2& Β≅δΓ (∃
−+∀.)%& Β]0ΑΓ8
Journal of Cell Science
4336
Journal of Cell Science 122 (23)
Fig. Ρ. Ξ:(ΚD 1&&(∗+1)+(∀ 5+)4 )4% WΧε ∃%<.+∃%& ΞΧΑ8 Β0εΓ!9+−%≅/1∋&% Η≅&%∃+%& −16+−.− ∋∃(?%∗)+(∀& (, ∗yo568∋93 ∗es67 ΒΓ &n, ∗yo568∋93 ∗es6! ΒεΓ ∗%//&
2.∃+∀3 −%+(&+&8 !−13%& 5%∃% ∗1∋).∃%2 %+)4%∃ %7%∃: Κ ΒΓ (∃ Μ ΒεΓ −+∀.)%&8 /)4(.34 Ξ:(ΚD ∃%−1+∀& /(∗1/+&%2 )( )4% WΧε )4∃(.34(.) −%+(&+& !! +∀ ∗es67 ∗%//& ΒΓ0
Ξ:(ΚD &+3∀1/ ∃1∋+2/: 2+&1∋∋%1∃& ,∃(− )4% WΧε (, ∗es6! ∗%//& 1∃∃%&)%2 1) )4% %∀2 (, −%+(&+& ! 1∀2 1&&(∗+1)%& 5+)4 )4% ∀.∗/%1∃ −%−=∃1∀% ΒεΓ8 ΒΡΓ!Ξ:(ΚD≅ςΑΧ
Β.∋∋%∃ ∋1∀%/Γ0 Ξ:(ΚD≅ςΑΧ Β−+22/% ∋1∀%/Γ 1∀2 Η∗Τ%2D≅Ξ:(ΚD Β/(5%∃ ∋1∀%/Γ /(∗1/+&% )( )4% WΧε& 1∀2 ∀.∗/%1∃ −%−=∃1∀% (, ∗es67 ∗%//& =%,(∃% 1∀1∋41&% !!8
ΒδΓ!∗yo568∋93 si,=8t,?o∗&to ∗es6! 3RΙϑD:c,c6Β,es:8ΕΓ≅ ∗%//& 5%∃% )∃%1)%2 5+)4 %+)4%∃ %)41∀(/ Β.∋∋%∃ ∋1∀%/&Γ (∃ %&)∃12+(/ Β/(5%∃ ∋1∀%/&Γ8 /)4(.34 %)41∀(/
41& ∀( %,,%∗) .∋(∀ )4% ∗%//./1∃ 2+&)∃+=.)+(∀ (, Ξ:(ΚD Β3∃%%∀0 1∃∃(54%12&Γ0 %&)∃12+(/≅+∀2.∗%2 Ρ2∗DΙ %6∋∃%&&+(∀ ∗1.&%2 Ξ:(ΚD )( ∃%).∃∀ )( )4% WΧε 1∀2 ∗(/(∗1/+&%
5+)4 )4% WΧε ∗(−∋(∀%∀) W+2Σ Β−13%∀)1Γ8
)4% 7+1=+/+): (, )4%&% ∗%//& Β91=/% DΓ8 94%&% 21)1 &.33%&) )41) Ξ:(ΚD
41& 1 ∃(/% +∀ −1+∀)1+∀+∀3 WΧε +∀)%3∃+): 2.∃+∀3 −%+(&+& 1∀2 )4%∃%=:
∗((∃2+∀1)+∀3 &∋(∃% ,(∃−1)+(∀ 5+)4 ∗4∃(−(&(−% &%3∃%31)+(∀ 2.∃+∀3
−%+(&+& !!8
Discussion
Myosin V function during conjugation
!∀ )4+& &).2:0 5% +∀7%&)+31)%2 )4% ,.∀∗)+(∀ (, )4% ,+&&+(∀ :%1&) −:(&+∀
α0 Ξ:(ΚD0 1∀2 ,(.∀2 )41) +) 41& &.=)/% ∃(/%& 2.∃+∀3 ∗(∀?.31)+(∀ 1∀2
+∀ )4% ∃%3./1)+(∀ (, &∋(∃% ,(∃−1)+(∀8 ⊥% 2+&∗(7%∃%2 )41) =()4 Ξ:(ΚD
1∀2 Ξ:(ΚΜ ∗(/(∗1/+&% 5+)4 1∗)+∀ )( )4% &+)% (, ∗%// ,.&+(∀ ΒΑ+38 ΦΓ8
94% 1∗)+∀ ∗:)(&>%/%)(∀ 41& 1∀ %&&%∀)+1/ ∃(/% 2.∃+∀3 ∗(∀?.31)+(∀0 1&
+//.&)∃1)%2 =: )4% ,1∗) )41) ∗%//& /1∗>+∀3 )4% −1)+∀3≅∗:∗/%≅&∋%∗+,+∗
,(∃−+∀ Α.&D0 1∃% &)%∃+/% ΒΧ%)%∃&%∀ %) 1/80 DΕΕΚΓ8 !∀ ∗(∀)∃1&) )( 5+/2≅
):∋% ∗%//&0 )4% ∗%// 51// 1) )4% &+)% (, ∗(∀?.31)+(∀ (, 133/.)+∀1)+∀3
9us6! ∗%//& ∃%−1+∀& +∀)1∗)0 ∋∃%7%∀)+∀3 )4% ,(∃−1)+(∀ (, 1 &+∀3/%
;:3()%8 !) +& 1∋∋%1/+∀3 )( ∗(∀∗/.2% )41) )4+& 9us6! ∋4%∀():∋% +&
=∃(.34) 1=(.) =: )4% 1=&%∀∗% (, Α.&D≅&%%2%2 1∗)+∀ ,+/1−%∀)&0 54+∗4
∋∃%7%∀)& Ξ:(ΚD 1∀2 Ξ:(ΚΜ ,∃(− 2%/+7%∃+∀3 ∗1∃3(%& ∃%<.+∃%2 ,(∃
∗%// ,.&+(∀ )( )4% &+)% (, ∗(∀?.31)+(∀8 Η(5%7%∃0 1& ∗%//& /1∗>+∀3 =()4
−:(&+∀ α ∋∃()%+∀& Β∗yo56! ∗yo5:!Γ 1∃% ∀() &)%∃+/% Β∀() &4(5∀Γ0
+) 5(./2 &.33%&) )4%&% )5( −()(∃& 1∃% ∀() )4% (∀/: ∋∃()%+∀& ∃%<.+∃%2
,(∃ ∃%∗∃.+)+∀3 ∗%//≅51//≅2%3∃12+∀3 %∀;:−%& )( )4% ∗%// )+∋8 94% ,1∗)
)41) (7%∃%6∋∃%&&+(∀ (, )4% Ξ:(ΚD ς9δ 1,,%∗)& ∋∃(∋%∃ ∗%// ,.&+(∀
&.33%&)& )41) +)& ∗1∃3(%& 1∃% =%+∀3 &%<.%&)%∃%2 =: )4+& ∗1∃3(≅=+∀2+∀3
)1+/ 2(−1+∀0 1∀2 ∋∃%7%∀)+∀3 )4%+∃ ∃%∗∃.+)−%∀) )( )4% &+)% (,
∗(∀?.31)+(∀0 =: 1∀ 1/)%∃∀1)+7% −%∗41∀+&− )41) +& +∀2%∋%∀2%∀) (,
−:(&+∀ α (∃ Α.&D ΒΡ1∃)13%∀1≅φ+∃(/1 %) 1/80 ΜΝΝ_Γ8
94% +2%∀)+): (, ∋()%∀)+1/ ∗1∃3(%& ∃%−1+∀& .∀>∀(5∀0 4(5%7%∃0 +)
+& +∀)∃+3.+∀3 )( &∋%∗./1)% (∀ )4%+∃ +2%∀)+):8 & −c/i0os&cc/&ro∗yces
3o∗4e ∗%//& 2( ∀() ∗(∀)1+∀ 1∀: (=7+(.& 4(−(/(3.%& (, >∀(5∀
%6(3/.∗1∀1&%& (∃ ∋∃(2.∗% 1∀: 2%)%∗)1=/% %6(3/.∗1∀1&% 1∗)+7+):
ΒΞ(/%∃( %) 1/80 DΕΕΕΓ0 ∗1∀2+21)%& ,(∃ ∗(∀?.31)+(∀≅&∋%∗+,+∗ −:(&+∀
α ∗1∃3(%& +∀∗/.2% )4% %∀2(3/.∗1∀1&%& ]∀3D Β1∀ %∀2(≅#≅D0Ι≅
3/.∗1∀1&%Γ0 3∀D 1∀2 3∀Μ Β%∀2(≅∃≅D0Ι≅3/.∗1∀1&%&Γ0 54+∗4 1∃%
∃%<.+∃%2 ,(∃ )4% 4:2∃(/:&+& (, ∗%// 51// &.31∃& 5+)4+∀ )4% ∋∃+−1∃:
&%∋).− 2.∃+∀3 ∗:)(>+∀%&+& Βδ%>>%∃ %) 1/80 ΜΝΝΣϑ Ξ1∃)+∀≅Ρ.12∃12(
%) 1/80 ΜΝΝΙΓ8 /)4(.34 1 ∃(/% ,(∃ 3∀Μ 41& =%%∀ .∀∗(7%∃%2 2.∃+∀3
)4% /1)%∃ &)13%& (, −%+(&+& Βδ%>>%∃ %) 1/80 ΜΝΝΥΓ0 )4% ,1∗) )41)
∗(−=+∀1)+(∀& (, &∋n60 en∋6 1∀2 &∋n: −.)1∀)& 1∃% ∀() &)%∃+/%
&.33%&)& )41) )4%: 1∃% ∀() ∃%<.+∃%2 ,(∃ ∗%// ,.&+(∀ )( (∗∗.∃8 !) 41&
=%%∀ &4(5∀ )41) )4% ,+&&+(∀ :%1&) %6(∗:&) ∗(−∋/%6 +& +∀7(/7%2 +∀
)1∃3%)+∀3 %∀;:−%&0 +∀∗/.2+∀3 %∀2(3/.∗1∀1&%& ∃%<.+∃%2 ,(∃ &%∋).−
∗/%1713% ΒΞ1∃)+∀≅Ρ.12∃12( %) 1/80 ΜΝΝΚϑ ⊥1∀3 %) 1/80 ΜΝΝΜΓ8 !) +&
)4%∃%,(∃% ∋(&&+=/% )41)0 1& +∀ =.22+∀3 :%1&)0 1 −:(&+∀ α ∋∃()%+∀
)1∃3%)& &%∗∃%)(∃: 7%&+∗/%& )( &∋%∗+,+∗ ∗%//./1∃ /(∗1)+(∀& ΒΧ∃.:∀% %)
1/80 DΕΕΦΓ0 54%∃% )4%: 1∗)+71)% %6(∗:&)≅2%∋%∀2%∀) %∀;:−%& 2.∃+∀3
∗%// ,.&+(∀8 Α.∃)4%∃ +∀7%&)+31)+(∀& 5+// 4(∋%,.//: .∀∗(7%∃ )4%
Myosin V and fission yeast meiosis
4337
−(/%∗./%Β&Γ )41) Ξ:(ΚD 1∀2 Ξ:(ΚΜ 2%/+7%∃ )( )4% ,.&+(∀ &+)%0 )(
1//(5 ∀(∃−1/ ∗(∀?.31)+(∀ )( (∗∗.∃8
Myosin V at microtubule-organising centres
Journal of Cell Science
W+−+/1∃/: )( ∗/1&& !! 1∀2 [ −:(&+∀&0 −:(&+∀ α& 417% =%%∀ ∃%∋(∃)%2
)( /(∗1/+&% )( )4% −+∗∃().=./%≅(∃31∀+&+∀3 ∗%∀)∃%& +∀ 1 ∀.−=%∃ (,
7%∃)%=∃1)% ∗%// ):∋%& Β]&∋∃%1,+∗( %) 1/80 DΕΕΦϑ 9&1>∃1>/+2%& %) 1/80
DΕΕΕΓ0 =.) 1/)4(.34 −:(&+∀ !! 1∀2 [ 1∃% >∀(5∀ )( =% +∀7(/7%2 +∀
)4% ∋(&+)+(∀+∀3 1∀2 1&&%−=/: (, )4% −%+()+∗ &∋+∀2/% ΒW∗4.4 1∀2
]//%∀=%∃30 ΜΝΝΦϑ ⊥%=%∃ %) 1/80 ΜΝΝΣΓ0 )4% ,.∀∗)+(∀ (, −:(&+∀ α1 1)
)4% Ξ9∴Ρ +& .∀>∀(5∀8 W+−+/1∃/: )( Ξ:(ΚD0 1,)%∃ ∃%∗∃.+)−%∀) )(
)4% Ξ9∴Ρ )4% /(∗1/+&1)+(∀ (, −:(&+∀ α1 +& 1∗)+∀ +∀2%∋%∀2%∀)
Β9&1>∃1>/+2%& %) 1/80 DΕΕΕΓ0 &.33%&)+∀3 )41) (∀∗% ∃%∗∃.+)%20 )4%+∃
/(∗1/+&1)+(∀ )( )4% Ξ9∴Ρ (∗∗.∃& +∀2%∋%∀2%∀)/: (, +)& −()(∃ 1∗)+7+):8
Χ∃%/+−+∀1∃: ΑΤΧ 21)1 &.33%&) )41) Ξ:(ΚD −()(∃ 1∗)+7+): +&
∃%<.+∃%2 )( ∃%∗∃.+) Ξ:(ΚD )( )4% WΧε Β∀() &4(5∀Γ8 94+& +& ∗(∀&+&)%∀)
5+)4 (.∃ (=&%∃71)+(∀ )41) )4% Ξ:(ΚD )1+/ 1/(∀% +& +∀&.,,+∗+%∀) ,(∃
WΧε 1&&(∗+1)+(∀0 1∀2 +∀ ∗(∀)∃1&) )( −.∃+∀% −:(&+∀ α10 %6∋∃%&&+(∀
(, Ξ:(ΚD≅ς9δ 1/(∀% +& +∀&.,,+∗+%∀) )( 2+&∋/1∗% )4% %∀2(3%∀(.&
,.//≅/%∀3)4 ∋∃()%+∀ ,∃(− )4% Ξ9∴Ρ +∀ ,+&&+(∀ :%1&) Β∀() &4(5∀Γ8
94%&% 21)1 +∀2+∗1)% )41) Ξ:(ΚD −()(∃ 1∗)+7+): +& ∃%<.+∃%2 ,(∃ +)&
∃%∗∃.+)−%∀) )( )4% WΧε0 54%∃% )4% )1+/ )4%∀ 1&&(∗+1)%& 5+)4 1
∗(−∋(∀%∀) (, )4% −%+()+∗ WΧε8
Myosin V and fungal meiosis
Fig. Σ. Ξ:(ΚD +& ∃%<.+∃%2 ,(∃ ∀(∃−1/ −%+()+∗ ∗4∃(−(&(−% &%3∃%31)+(∀ 1∀2
&∋(∃% ,(∃−1)+(∀8 Β0εΓ!Χ41&%≅∗(∀)∃1&) +−13%& Β/%,) ∋1∀%/&Γ 1∀2 δΧ! &)1+∀+∀3
Β∃+34) ∋1∀%/&Γ (, 1&∗+ ∋∃(2.∗%2 ,∃(− ∗∃(&&%& =%)5%%∀ ∗yo567 ΒΓ (∃ ∗yo56!
∗%//& ΒεΓ8 ΒΡΓ!ςΑΧ &+3∀1/ Β/%,) ∋1∀%/Γ −%∃3%2 5+)4 ∋41&%≅∗(∀)∃1&) +−13% Β∃+34)
∋1∀%/Γ ,∃(− ,%c:8∋93 ∗yo56! ∗%//& +//.&)∃1)% ∀.∗/%1∃ −+&≅&%3∃%31)+(∀ 2%,%∗)& +∀
)4%&% ∗%//&8 Βδ0]Γ!W+2Σ≅)(−1)( Β−13%∀)1Γ 1∀2 δ/∗Μ Β3∃%%∀Γ &+3∀1/& ∃%7%1/ WΧε
2%,%∗)& +∀ ∗yo56! ∗%//&0 54+∗4 (,)%∀ /%12 )( 2+&∃.∋)+(∀ (, ∀.∗/%1∃ −%−=∃1∀%
+∀)%3∃+): Β1∃∃(54%12&Γ8 ΒΑ0ςΓ!∴7%∃%6∋∃%&&+(∀ (, ,.//≅/%∀3)4 Ξ:(ΚD /%12& )( )4%
,(∃−1)+(∀ (, 1&∗+ ∗(∀)1+∀+∀3 1∀ +∀1∋∋∃(∋∃+1)% ∀.−=%∃ (, &∋(∃%&8 ΒΑΓ!94%&%
&∋(∃%& (,)%∀ ∗(∀)1+∀%2 1=%∃∃1∀) WΧε −1)%∃+1/ ΒW+2Σ0 −13%∀)1Γ0 ∃1)4%∃ )41∀
∀.∗/%+ Βδ/∗Μ0 3∃%%∀Γ8 W∗1/% =1∃&β Κ!∀−8
/)4(.34 ∗/1&& α −:(&+∀& 417% ∀() =%%∀ &%%∀ )( /(∗1/+&% )( −%+()+∗
WΧε& +∀ ()4%∃ ,.∀3+0 )4%&% −()(∃& 417% =%%∀ +−∋/+∗1)%2 +∀
)∃1∀&∋(∃)+∀3 ∋∃%&∋(∃% −%−=∃1∀% ∋∃%∗.∃&(∃& )( )4% &+)% (, &∋(∃%
,(∃−1)+(∀ 1∀2 ∗(∀?.31)+(∀ ).=% ,(∃−1)+(∀ +∀ =.22+∀3 :%1&) 1∀2 )4%
∋/1∀) ∋1)4(3%∀0 Usti%&∋o ∗&y,is Β916+& %) 1/80 ΜΝΝ_ϑ ⊥%=%∃ %) 1/80
ΜΝΝΙΓ0 &.33%&)+∀3 1 ∗(∀&%∃7%2 −%+()+∗ ,.∀∗)+(∀ ,(∃ )4+& ∗/1&& (,
−()(∃ ∋∃()%+∀& +∀ ,.∀3+8 94% ,+&&+(∀ :%1&) WΧε .∀2%∃3(%& &)∃.∗).∃1/
∗41∀3%& 2.∃+∀3 −%+(&+& ΒΗ+∃1)1 1∀2 W4+−(210 DΕΕΣϑ 91∀1>1 1∀2
Η+∃1)10 DΕΦΜΓ 54%∀ +) 2%7%/(∋& +∀)( 1 &+)% ,(∃ ,(∃%&∋(∃% −%−=∃1∀%
(∃31∀+&1)+(∀8 ⊥% 2+&∗(7%∃%2 )41) Ξ:(ΚD ∃%−1+∀& 1&&(∗+1)%2 5+)4
)4% ∀.∗/%1∃ −%−=∃1∀% .∀)+/ (∀&%) (, −%+(&+& !!0 54%∀ −(&) (, )4%&%
&)∃.∗).∃1/ ∗41∀3%& (∗∗.∃ )( )4% (.)%∃ ∋/1<.% (, )4% WΧε ΒW4+−(210
ΜΝΝΣΓ8 94+& &.33%&)& )4% %6+&)%∀∗% (, 1 ∗(∀)∃(/ −%∗41∀+&− )(
∃%3./1)% Ξ:(ΚD ∃%∗∃.+)−%∀) )( )4% WΧε 1∀2 )4% )+−+∀3 (, ,(∃% &∋(∃%
−%−=∃1∀% 1&&%−=/:8 94+& +& ∀() (∀/: ∗(∀&+&)%∀) 5+)4 )4% ∃%2.∗)+(∀
+∀ 7+1=+/+): (, &∋(∃%& ∋∃(2.∗%2 ,∃(− 1 ∗∃(&& =%)5%%∀ ∗%//& /1∗>+∀3
Ξ:(ΚD0 =.) +& 1/&( &.∋∋(∃)%2 =: (.∃ .∀∋.=/+&4%2 ,+∀2+∀3 )41) Ξ:(ΚD
+∀)%∃1∗)& 5+)4 )4% ∗4%∗>∋(+∀) ∋∃()%+∀ Τ12ΜΣ8 94% ,.∀∗)+(∀ (, )4+&
∋∃()%+∀ +& %&&%∀)+1/ 2.∃+∀3 1 2+∋/(+2 /+,%∗:∗/% Β91∀1>1 %) 1/80 ΜΝΝΝΓ0
=%∗1.&% +) 1∗)& 1& 1 ∃%3./1)(∃ (, ε:∃Μ0 1 ∗(−∋(∀%∀) (, )4% &+3∀1//+∀3
∋1)451: )41) +& %&&%∀)+1/ ,(∃ =()4 ∗(∀?.31)+(∀ 1∀2 −%+(&+& ΒΛ?1%∃./,,
%) 1/80 ΜΝΝΚϑ ∴;(% %) 1/80 ΜΝΝΜϑ ⊥1∀3 %) 1/80 DΕΕDΓ8 Η(5%7%∃0 )4%
−(&) 2∃1−1)+∗ 1∀2 ∗(−∋%//+∀3 %7+2%∀∗% )( &.∋∋(∃) )4% %6+&)%∀∗% (,
1 Ξ:(ΚD≅2%∋%∀2%∀) −%∗41∀+&− )( −1+∀)1+∀ WΧε +∀)%3∃+): 2.∃+∀3
−%+(&+& !! 1) )4% (∀&%) (, ,(∃% &∋(∃% −%−=∃1∀% 1&&%−=/: ∗(−%&
,∃(− )4% ,+∀2+∀3 )41) (7%∃%6∋∃%&&+(∀ (, )4% ,.//≅/%∀3)4 Ξ:(ΚD
Table 1. Percentage frequency of defects in spore formation and viability upon modulation of Myo51 expressionς
∗yo567 " ∗yo567
∗yo567" ∗yo56!
∗yo56! " ∗yo56!
Ξ:(ΚD (7%∃%6∋∃%&&+(∀
Σ &∋(∃%&χΣ ∀.∗/%+
Σ &∋(∃%&χΙ ∀.∗/%+
λΣ &∋(∃%&
W∋(∃% 7+1=+/+):
ΕΚ8ΥΝmΜ8ΥΚ
ΕD8ΦΝmD8ΦΕ
ΦΥ8ΙΝmD8ΚΝ
_Ε8ΚΝm_8ΙΕ
Ι8DΥmD8ΣΣ
Κ8ΦΙmΜ8ΜΚ
Ε8DΥmΜ8ΝΜ
Φ8ΥΚm_8ΝD
D8DΥmD8_
Μ8ΙΝmΝ8Υ_
Ι8ΚΝmΜ8ΜΕ
ΜD8ΥΚmΝ8ΙΚ
ΕΙ8ΜmΚ8Μ
ΕD8Μm_8Υ
_Σ8DmΦ8Ι
Σ_8DmΙ8Υ
νδ1)1 17%∃13%2 ,∃(− )4∃%% +∀2%∋%∀2%∀) %6∋%∃+−%∀)&8
4338
Journal of Cell Science 122 (23)
Journal of Cell Science
Fig. 8. W∗4%−1)+∗ ∃%∋∃%&%∀)1)+(∀ (, /(∗1/+&1)+(∀ (, Ξ:(ΚD 1∀2
1∗)+∀ )4∃(.34(.) )4% ,+&&+(∀ :%1&) −%+()+∗ /+,%∗:∗/%8
∋∃()%+∀ ∗1∀ =∃+∀3 1=(.) )4% ,(∃−1)+(∀ (, 1&∗+ ∗(∀)1+∀+∀3 1 4.3%
∀.−=%∃ (, &∋(∃%&0 )4% −1?(∃+): (, 54+∗4 1∃% .∀7+1=/%8
MPF-dependent regulation of the association of Myo51 with
the SPB
/)4(.34 Ξ:(ΚD −+34) 417% 1 ∃(/% −1+∀)1+∀+∀3 )4% WΧε 2.∃+∀3
−%+(&+&0 +) +& .∀∗/%1∃ 4(5 +)& (5∀ ∃%∗∃.+)−%∀) +& ∃%3./1)%28 ΑΤΧ
1∀1/:&+& (, )4% Ξ:(ΚD WΧε &+3∀1/ &.33%&)& )41) Ξ:(ΚD ).∃∀(7%∃
1) )4% WΧε +& ∀(∃−1//: ∃%2.∗%2 1& −%+(&+& ∋∃(3∃%&&%&8 Ψ() (∀/: +&
)4+& ∗41∀3% +∀ ).∃∀(7%∃ 1 ∃%,/%∗)+(∀ (, )4% ∃%2.∗)+(∀ +∀ ∗%//./1∃ /%7%/&
(, Ρ2∗DΙ ΒΡ:∗/+∀εΓ0 =.) 1/&(0 54%∀ Ξ%&D +& ∀() ∋∃%&%∀) +∀ )4% ∗%//
)( ∋∃%7%∀) ∋∃%−1).∃% 2%3∃121)+(∀ (, )4% ∋((/ (, ∗:∗/+∀ ε ∃%<.+∃%2
,(∃ −%+(&+& !! (∀&%) Β!;151 %) 1/80 ΜΝΝΚΓ0 )4% Ξ:(ΚD &+3∀1/ ∃1∋+2/:
2+&1∋∋%1∃& ,∃(− )4% WΧε8 94% ,1∗) )41) +) 51& ∋(&&+=/% )( ∗(.∀)%∃1∗)
)4+& %,,%∗) )4∃(.34 )4% ∃1∋+2 +∀2.∗)+(∀ (, ∀(∀≅2%3∃121=/% Ρ2∗DΙ
+∀2+∗1)%& )4% Ρ2∗Μ≅∗:∗/+∀≅ε ∗(−∋/%6 +& ∃%<.+∃%2 ,(∃ /(∗1/+&1)+(∀ (,
Ξ:(ΚD 1) )4% −%+()+∗ Ξ9∴Ρ&8 !∀ 122+)+(∀ )( )4+& ,+∀2+∀3 )41) ΞΧΑ
∃%3./1)%& Ξ:(ΚD −%+()+∗ /(∗1/+&1)+(∀0 Ρ2∗Μ 41& =%%∀ &4(5∀ )( 417%
1∀ +−∋(∃)1∀) ∃(/% +∀ ∃%3./1)+∀3 −%+(&+& !! ∋∃(3∃%&&+(∀0 =.) &+−+/1∃/:
)( Ξ:(ΚD0 +) 1/&( ∗((∃2+∀1)%& )4+& 5+)4 &.=&%<.%∀) &∋(∃% ,(∃−1)+(∀
Βδ+&∗4+∀3%∃ %) 1/80 ΜΝΝΦϑ Η1:/%& %) 1/80 DΕΦ_Γ8
!) +& .∀∗/%1∃ 54: ∀(∃−1/ &∋+∀2/% ,(∃−1)+(∀ 2+2 ∀() (∗∗.∃ .∋(∀
+∀2.∗)+(∀ (, ∀(∀≅2%3∃121=/% Ρ2∗DΙ +∀ 1∃∃%&)%2 ∗es6! ∗%//&8 !) ∗(./2
∋%∃41∋& +∀2+∗1)% )41) Ξ%&D ∋∃%7%∀)& 2%3∃121)+(∀ (, &.=≅∋(∋./1)+(∀&
(, ()4%∃ 1& :%) .∀2%)%∃−+∀%2 ∋∃()%+∀& ∃%<.+∃%2 ,(∃ −%+(&+& !! &∋+∀2/%
,(∃−1)+(∀0 (∃ )41) Ξ%&D +& ∃%<.+∃%2 ,(∃ ,.∃)4%∃ .∀%6∋/(∃%2 ,.∀∗)+(∀&
2.∃+∀3 −%+(&+&8 Η(5%7%∃0 %6∋∃%&&+(∀ (, )4% ∀(∀≅2%3∃121=/% ∗:∗/+∀
ε 51& &.,,+∗+%∀) ,(∃ %∀(.34 1∗)+71)+(∀ 2.∃+∀3 −%+(&+& !! )( ∋∃(−()%
)4% Ξ:(ΚD≅WΧε 1&&(∗+1)+(∀ 1∀2 ∋∃(7+2%& 1∀ +∀&+34) +∀)( )4%
∃%3./1)+(∀ 1∀2 ,.∀∗)+(∀ (, )4+& 1∗)+∀≅1&&(∗+1)%2 −()(∃8
Does Myo51 always deliver the goods?
Ρ/1&& α −:(&+∀& 5+)4+∀ 7%∃)%=∃1)% ∗%//& %6+&) 1& 2+−%∃+∗ ∋∃(∗%&&+7%
−()(∃& )( )∃1∀&∋(∃) ∗%//./1∃ ∗1∃3(%& )4∃(.34(.) )4% ∗%// ΒΞ%4)1 %)
1/80 DΕΕΕϑ Ξ%∃−1// %) 1/80 DΕΕΦΓ8 /)4(.34 −:(&+∀ α ∋∃()%+∀& ,∃(−
&+−∋/%∃ (∃31∀+&−& 2%−(∀&)∃1)% ∀(∀≅∋∃(∗%&&+7% +∀ 7+)∃( −()(∃
1∗)+7+): ΒΛ∃%−%∀)&(71 %) 1/80 ΜΝΝ_ϑ Τ%∗>≅Χ%)%∃&(∀ %) 1/80 ΜΝΝDϑ 9()4
%) 1/80 ΜΝΝΚΓ0 +) +& ∗/%1∃ )41) )4%: &)+// ∃%)1+∀ )4% 1=+/+): )( −(7% ∃1∋+2/:
)4∃(.34(.) )4% ∗%// Βς∃1//%∃) %) 1/80 ΜΝΝΥϑ W∗4()) %) 1/80 ΜΝΝΜΓ8
/)4(.34 )4% ∋∃(∗%&&+7+): (, −1−−1/+1∀ −:(&+∀ α 1∀2 >+∀%&+∀&
∗1∀ =% %∀41∀∗%2 =: 1&&(∗+1)+∀3 5+)4 %1∗4 ()4%∃ Β/+ %) 1/80 ΜΝΝΦΓ0
)4% ∋∃(∗%&&+7+): (, )4% =.22+∀3 :%1&) ∗/1&& α −:(&+∀ Ξ:(Σ0 +&
∃%3./1)%2 =: =+∀2+∀3 ∋1∃)∀%∃& )41) ∃%3./1)% +)& 1=+/+): )( 2+−%∃+&%0
1∀2 )4%∃%=: 51/> 1/(∀3 1∗)+∀ ΒΗ(23%& %) 1/80 ΜΝΝΦΓ8 94% 21)1
∋∃%&%∀)%2 4%∃% &.33%&) )41) 1 &+−+/1∃ −%∗41∀+&− −+34) %6+&) +∀
,+&&+(∀ :%1&) )( ∃%3./1)% Ξ:(ΚD0 )4% 1∀1/(3.% (, Ξ:(Σ +∀ =.22+∀3
:%1&)8
/)4(.34 +) +& ∗/%1∃ )41) Ξ:(ΚD +& ∃%<.+∃%2 ,(∃ ∋∃(∋%∃ ∗%// ,.&+(∀
2.∃+∀3 ∗(∀?.31)+(∀0 1∀2 +) +& ∋(&&+=/% )( (=&%∃7% Ξ:(ΚD −(7+∀3
.∋(∀ 1∗)+∀ ,+/1−%∀)& )( )4% &+)% (, ∗%// ,.&+(∀ Β∀() &4(5∀Γ0 +) +& ∀()
∗/%1∃ 54%)4%∃ Ξ:(ΚD +& ∃%<.+∃%2 )( 2%/+7%∃ ∗1∃3(%& )( )4% −%+()+∗
WΧε8 ⊥% 417%0 1& :%)0 =%%∀ .∀1=/% )( 2%)%∗) Ξ:(ΚD ,(∗+ −(7+∀3
)( (∃ ,∃(− )4% −%+()+∗ WΧε8 94%&% 21)1 +//.&)∃1)% )41) )4%∃% +& 1 /1∃3%
∀(∀≅2:∀1−+∗ ∋(∋./1)+(∀ (, Ξ:(ΚD 1) )4+& (∃31∀%//% )4∃(.34(.)
−%+(&+& !!0 1∀2 1∗)+∀ +& ∀() ∃%<.+∃%2 ,(∃ Ξ:(ΚD )( −1+∀)1+∀ +)&
∋(&+)+(∀ )4%∃%8 Ξ:(ΚD )4%∃%,(∃% ∋∃(=1=/: +∀)%∃1∗)& 5+)4 )4% WΧε
7+1 +)& )1+/ 2(−1+∀ 2.∃+∀3 )4+& ∋%∃+(20 54+∗4 %6∋/1+∀& +)& 1=+/+): )(
/(∗1/+&% )4%∃% +∀ )4% 1=&%∀∗% (, 1∗)+∀8 94+& /%17%& +)& −()(∃ 2(−1+∀
,∃%% )( +∀)%∃1∗) 5+)4 1∗)+∀0 1∀2 )4%∃%=: 1∗) 1& 1∀ 1∀∗4(∃ (∃ )%)4%∃
,(∃ )4% 3∃(5+∀3 1∗)+∀ ∋(/:−%∃& )( ,1∗+/+)1)% ,(∃%&∋(∃% −%−=∃1∀%
,(∃−1)+(∀ ΒΧ%)%∃&%∀ %) 1/80 DΕΕΦ=ϑ 916+& %) 1/80 ΜΝΝ_Γ8 94.&0 Ξ:(ΚD
−+34) ∀() (∀/: 417% )4% 1=+/+): )( 2%/+7%∃ ∗1∃3(%& )( &∋%∗+,+∗ ∗%//./1∃
/(∗1)+(∀&0 =.) ∗(./2 1/&( 1∗) 1& 1 )%)4%∃ (∃ )%∀&+(∀ &%∀&(∃ 5+)4+∀ )4%
∗%//0 +∀ 1 ∗%//≅∗:∗/%≅ 1∀2 +∀2%%2 /+,%∗:∗/%≅2%∋%∀2%∀) −1∀∀%∃8
Materials and Methods
Yeast cell culture and strains
−Κ 3o∗4e &)∃1+∀& .&%2 +∀ )4+& &).2: 1∃% /+&)%2 +∀ &.∋∋/%−%∀)1∃: −1)%∃+1/ 91=/% WD8
Ρ%// ∗./).∃% 1∀2 −1+∀)%∀1∀∗% 5%∃% ∗1∃∃+%2 (.) 1& 2%&∗∃+=%2 ΒΞ(∃%∀( %) 1/80 DΕΕDΓ8
Ρ%//& 5%∃% 3∃(5∀ +∀ &.∋∋/%−%∀)%2 −+∀+−1/ −%2+.− Β]ΞΞΜΓ (∃ &∋(∃./1)+(∀ −%2+.−
ΒΞWφΓ Β]3%/ %) 1/80 DΕΕΣΓ8
Molecular biology
W)∃1+∀& %6∋∃%&&+∀3 ,/.(∃(∋4(∃%≅)133%2 Ξ:(ΚD 1) %+)4%∃ +)& Ψ≅)%∃−+∀.& Β∗yo568n>Λϑ
1∀2 ∗yo568nΣΕcRe,6Γ (∃ Ρ≅)%∃−+∀.& Β∗yo568∗Χ/erryΓ 5%∃% ∗∃%1)%2 .&+∀3 )5(
2+&)+∀∗) −%)4(2&8 94% ∗yo568∗Χ/erry 1//%/%0 +∀ 54+∗4 )4% Ι# %∀2 (, )4% %∀2(3%∀(.&
∗4∃(−(&(−1/ ∗(∋: (, )4% ∗yo567 ∋ene 51& ,.&%2 )( ∗δΨ %∀∗(2+∀3 )4% −Ρ4%∃∃:
,/.(∃(∋4(∃% ΒW41∀%∃ %) 1/80 ΜΝΝΣΓ 51& ∗∃%1)%2 .&+∀3 1 −%)4(2 2%&∗∃+=%2 ∋∃%7+(.&/:
Myosin V and fission yeast meiosis
Βε14/%∃ %) 1/80 DΕΕΦΓ .&+∀3 ∋Α_1≅∗4%∃∃:≅>1∀Ξ[_ 1& 1 )%−∋/1)% Β91∀1>1 %) 1/80 ΜΝΝΚΓ8
W)∃1+∀& %6∋∃%&&+∀3 Ψ≅)%∃−+∀1/ ,.&+(∀ ∋∃()%+∀& 5%∃% ∗∃%1)%2 =: +&(/1)+∀3 )4% ∗yo567
3%∀% 1& 1 −&%!≅Γ&∗Η! ,∃13−%∀) ,∃(− ∋Τ]ΧΣD∗yo567 Β⊥+∀ %) 1/80 ΜΝΝDΓ 1∀2 ∗/(∀+∀3
)4+& +∀)( )4% −&%!≅Γ&∗Η! &+)%& (, )4% ∋/1&−+2& ∋!Ψ9ΣDΨςΑΧ 1∀2 ∋!Ψ9ΣDΨΣΗ∗Τ%2D
)( ∗∃%1)% ∋!Ψ9ΣDΨςΑΧ∗yo56ο 1∀2 ∋!Ψ9ΣDΨΣΗ∗Τ%2D∗yo5678 94%&% ∋/1&−+2& 5%∃%
/+∀%1∃+&%2 1∀2 +∀)%3∃1)%2 +∀)( )4% %eu67 /(∗.& )( ∗∃%1)% )4% ∗yo568n>Λϑ 1∀2 ∗yo568
n=ΕcRe,6 1//%/%&0 ∃%&∋%∗)+7%/:8 ∀ &ct68n>Λϑ &)∃1+∀ 51& ∗∃%1)%2 =: +&(/1)+∀3 )4%
&ct67 ∗δΨ 1& 1 −&%! ,∃13−%∀) =: ΧΡΤ0 1∀2 ∗/(∀+∀3 )4+& +∀)( ∋!Ψ9ΦDΨςΑΧ0 54+∗4
51& &.=&%<.%∀)/: &%<.%∀∗%20 /+∀%1∃+&%2 1∀2 +∀)%3∃1)%2 +∀)( )4% %eu67 /(∗.&8 94%
∗yo568>?≅n>Λϑ 1//%/%0 %∀∗(2+∀3 )4% Ξ:(ΚD 3/(=./1∃ )1+/ 2(−1+∀ ,.&%2 )( ςΑΧ 1)
+)& Ψ≅)%∃−+∀.&0 51& ∗∃%1)%2 =: +&(/1)+∀3 δΨ %∀∗(2+∀3 ∃%&+2.%& ΕΕ_≅DΣΥD (, ∗yo567
1& 1 −&%!≅Γ&∗Η! ,∃13−%∀) =: ΧΡΤ0 54+∗4 51& &%<.%∀∗%2 1∀2 ∗/(∀%2 +∀)( )4% −&%!≅
Γ&∗Η! &+)%& (, ∋!Ψ9ΣDΨςΑΧ )( ∗∃%1)% ∋!Ψ9ΣDΨςΑΧ−:(ΚD≅ς9δ8 94+& 51&
&.=&%<.%∀)/: /+∀%1∃+&%2 1∀2 +∀)%3∃1)%2 +∀)( )4% %eu67 /(∗.&8
Journal of Cell Science
Fluorescence microscopy
Ρ%//& 5%∃% −(.∀)%2 +∀ 1 ε+(∋)%∗4& ΑΡWΜ Βε+(∋)%∗4&0 ε.)/%∃0 ΧΓ 1∀2 )4% &1−∋/% 4(/2%∃0
(=?%∗)+7% /%∀& 1∀2 %∀7+∃(∀−%∀)1/ ∗41−=%∃ 5%∃% −1+∀)1+∀%2 1) )4% ∃%<.+∃%2 )%−∋%∃1).∃%8
W1−∋/%& 5%∃% 7+&.1/+&%2 .&+∀3 (∀% (, )5( &:&)%−&8 94% ,+∃&) 51& =1&%2 .∋(∀ 1∀
∴/:−∋.& ![ΥD −+∗∃(&∗(∋% 5+)4 1 Χ/1∀∋( "DΝΝ 9!ΤΑΞ≅WΧ D8ΣΚ Ψ /%∀& −(.∀)%2
(∀ 1 Χ!Α∴Ρ Η≅16+& ,(∗.& 2∃+7% ΒΧ4:&+> !∀&)∃.−%∀)&0 Λ1∃/&∃.4%0 ς%∃−1∀:Γ0 1∀2
+//.−+∀1)%2 5+)4 1∀ 1.)(−1)%2 ΙΝΝ ⊥ [%∀(∀ /+34) &(.∃∗% ΒW.))%∃0 Ψ(71)(0 ΡΓ .&+∀3
1∋∋∃(∋∃+1)% ,+/)%∃& 1∀2 7+&.1/+&%2 .&+∀3 %+)4%∃ 1 Ρ((/&∀1∋ Ηπ (∃ π.1∀)]Ξ ΡΡδ ∗1−%∃1
ΒΧ4()(−%)∃+∗&0 9.∗&(∀0 γΓ ∗(∀)∃(//%2 5+)4 Ξ%)1−(∃∋4 &(,)51∃% ΒΞ(/%∗./1∃ δ%7+∗%&0
δ(5∀+∀3)(∀0 ΧΓ8 94% &%∗(∀2 &:&)%− 51& 1 δ%/)17+&+(∀ Ρ(∃% Β∋∋/+%2 Χ∃%∗+&+(∀
!∀&)∃.−%∀)&Γ0 −(.∀)%2 (∀ 1∀ ∴/:−∋.& ![≅ΥD −+∗∃(&∗(∋% 54+∗4 .)+/+&%& 1∀ ∴/:−∋.&
Χ/1∀≅1∋(∗4∃(−1) "DΝΝ D8ΣΚ Ψ (=?%∗)+7% /%∀& 1∀2 7+&.1/+&1)+(∀ 7+1 1 Ρ1&∗12% !! ΚDΜε
]ΞΡΡδ ∗1−%∃1 ΒΧ4()(−%)∃+∗&Γ8 94+& &:&)%− 51& .&%2 )( 7+&.1/+&% =()4 %ςΑΧ 1∀2
29(−1)( ,/.(∃(∋4(∃%& .&+∀3 1 ΙΝΝ ⊥ [%∀(∀ Χ! /+34) &(.∃∗% 1∀2 ]9≅&%21) ,+/)%∃&
ΒΡ4∃(−1Γ8 94+& &:&)%− 41& 1∀ 122+)+(∀1/ ΚΝ −⊥ ΣΦΦ ∀− /1&%∃ ,(∃ ∋4()(=/%1∗4+∀3
+∀7%&)+31)+(∀& 1∀2 )4% /1&%∃ 1∋%∃).∃% 51& ∗1/+=∃1)%2 5+)4 1 &∋() &+;% (, Ν8Σ ∀−8
⊥% )41∀> Ο∃&./1 Α/%+30 ε%θ)1 ς∃1//%∃)0 !1+∀ Η131∀0 Λ1:(>( 91∀1>1
1∀2 )4% η1∋1∀ Ψ1)+(∀1/ Ζ%1&) ε+(≅∃%&(.∃∗% .∀+) ,(∃ &)∃1+∀& 1∀2 ∋/1&−+2&8
⊥% )41∀> Λ17+/ 9.∃ 1∀2 η(∀ Ψ%&& ,(∃ )%∗4∀+∗1/ 1&&+&)1∀∗% 1∀2 ∃−%∀+.&
Ρ%∀).∃+(∀0 3∀%& ς∃1//%∃) 1∀2 Ξ+>% ς%%7%& ,(∃ 2+&∗.&&+(∀& 1∀2
∗(−−%∀)& (∀ )4% −1∀.&∗∃+∋)8 94+& 5(∃> 51& &.∋∋(∃)%2 =: )4% εεWΤΡ0
Ρ1∀∗%∃ Τ%&%1∃∗4 ΟΛ 1∀2 )4% Τ1−(∀ ∃%∗%& ,(.∀21)+(∀8
References
Ali, M. Ξ., Lu, H., Bookwalter, C. S., Οarshaw, D. M. and Trybus, [. M. ΒΜΝΝΦΓ8
Ξ:(&+∀ α 1∀2 Λ+∀%&+∀ 1∗) 1& )%)4%∃& )( %∀41∀∗% %1∗4 ()4%∃&Θ ∋∃(∗%&&+7+):8 ϑrocΚ Μ&t%Κ
Νc&,Κ −ciΚ U−Ν 1050 Σ_ΕD≅Σ_Ε_8
Bahler, J., Οu, J. ⊥., Longtine, M. S., Shah, N. G., Mc[enzie, A., 3rd, Steever, A. B.,
Οach, A., Philippsen, P. and Pringle, J. R. ΒDΕΕΦΓ8 Η%)%∃(/(3(.& −(2./%& ,(∃ %,,+∗+%∀)
1∀2 7%∃&1)+/% ΧΡΤ≅=1&%2 3%∀% )1∃3%)+∀3 +∀ W∗4+;(&1∗∗41∃(−:∗%& ∋(−=%8 Οe&st 140 ΕΣΙ≅
ΕΚD8
Boe, C. A., Garcia, I., Pai, C. C., Sharom, J. R., Skjolberg, H. C., Boye, E., [earsey,
S., Macneill, S. A., Tyers, M. D. and Grallert, B. ΒΜΝΝΦΓ8 Τ1∋+2 ∃%3./1)+(∀ (, ∋∃()%+∀
1∗)+7+): +∀ ,+&&+(∀ :%1&)8 ΓΠΧ Χe%% Γio%Κ χ0 ΜΙ8
Cartagena-Lirola, H., Duran, A. and &aldivieso, M. H. ΒΜΝΝ_Γ8 94% W∗4+;(&1∗∗41∃(−:∗%&
∋(−=% ∗,∃Dο 3%∀% ∋1∃)+∗+∋1)%& +∀ −1)+∀3 )4∃(.34 1 ∀%5 ∋1)451: )41) +& +∀2%∋%∀2%∀) (,
,.&Dο8 Οe&st 230 ΙΥΚ≅ΙΦΦ8
Chikashige, Ξ., Ding, D. ⊥., Funabiki, H., Haraguchi, T., Mashiko, S., Ξanagida, M.
and Hiraoka, Ξ. ΒDΕΕΣΓ8 9%/(−%∃%≅/%2 ∋∃%−%+()+∗ ∗4∃(−(&(−% −(7%−%∀) +∀ ,+&&+(∀
:%1&)8 −cience 2Ρ40 ΜΥΝ≅ΜΥΙ8
Dekker, N., Speijer, D., Grun, C. H., van den Berg, M., de Haan, A. and Hochstenbach,
F. ΒΜΝΝΣΓ8 Τ(/% (, )4% 1/∋41≅3/.∗1∀1&% 3∀D∋ +∀ ,+&&+(∀≅:%1&) ∗%// &%∋1∃1)+(∀8 Πo%Κ Γio%Κ
Χe%% 150 ΙΕΝΙ≅ΙΕDΣ8
Dekker, N., van Rijssel, J., Distel, B. and Hochstenbach, F. ΒΜΝΝΥΓ8 Τ(/% (, )4% 1/∋41≅
3/.∗1∀1&% 3∀Μ∋ +∀ 1&∗.&≅51// %∀2(/:&+& ,(//(5+∀3 &∋(∃./1)+(∀ +∀ ,+&&+(∀ :%1&)8 Οe&st
240 ΜΥΕ≅ΜΦΦ8
Dischinger, S., [rapp, A., δie, L., Paulson, J. R. and Simanis, &. ΒΜΝΝΦΓ8 Ρ4%−+∗1/
3%∀%)+∗ 1∀1/:&+& (, )4% ∃%3./1)(∃: ∃(/% (, Ρ2∗Μ∋ +∀ )4% W8 ∋(−=% &%∋)1)+(∀ +∀+)+1)+(∀
∀%)5(∃>8 ΘΚ Χe%% −ciΚ 1210 ΦΣΙ≅ΦΚΙ8
Egel, R., Οiller, M., [jaerulff, S., Davey, J. and Nielsen, O. ΒDΕΕΣΓ8 &&%&&−%∀) (,
∋4%∃(−(∀% ∋∃(2.∗)+(∀ 1∀2 ∃%&∋(∀&% +∀ ,+&&+(∀ :%1&) =: 1 41/( )%&) (, +∀2.∗%2 &∋(∃./1)+(∀8
Οe&st 100 DΙΣΥ≅DΙΚΣ8
Espreafico, E. M., Coling, D. E., Tsakraklides, &., [rogh, [., Οolenski, J. S., [alinec,
G. and [achar, B. ΒDΕΕΦΓ8 φ(∗1/+;1)+(∀ (, −:(&+∀≅α +∀ )4% ∗%∀)∃(&(−%8 ϑrocΚ Μ&t%Κ
Νc&,Κ −ciΚ U−Ν χ50 Φ_Ι_≅Φ_ΣD8
Flory, M. R., Morphew, M., Joseph, J. D., Means, A. R. and Davis, T. N. ΒΜΝΝΜΓ8 Χ∗∋D∋0
1∀ W∋∗DDΝ∋≅∃%/1)%2 ∗1/−(2./+∀ )1∃3%) 1) )4% ∗%∀)∃(&(−% (, )4% ,+&&+(∀ :%1&)
W∗4+;(&1∗∗41∃(−:∗%& ∋(−=%8 Χe%% >roΡt/ ≅i99erΚ 130 ΣΥ≅ΚΦ8
Grallert, A., Martin-Garcia, R., Bagley, S. and Mulvihill, D. P. ΒΜΝΝΥΓ8 !∀ 7+7( −(7%−%∀)
(, )4% ):∋% α −:(&+∀ Ξ:(ΚΜ ∃%<.+∃%& 2+−%∃+&1)+(∀ =.) +& +∀2%∋%∀2%∀) (, )4% ∀%∗> 2(−1+∀8
ΘΚ Χe%% −ciΚ 1200 ΣΝΕΙ≅ΣΝΕΦ8
Hagan, I. and Ξanagida, M. ΒDΕΕΜΓ8 Λ+∀%&+∀≅∃%/1)%2 ∗.)Υ ∋∃()%+∀ 1&&(∗+1)%& 5+)4 −+)()+∗
1∀2 −%+()+∗ &∋+∀2/%& +∀ ,+&&+(∀ :%1&)8 Μ&ture 35Ρ0 ΥΣ≅Υ_8
4339
Hayles, J., Aves, S. and Nurse, P. ΒDΕΦ_Γ8 &.∗D +& 1∀ %&&%∀)+1/ 3%∀% +∀7(/7%2 +∀ =()4 )4%
∗%// ∗:∗/% 1∀2 3∃(5)4 +∀ ,+&&+(∀ :%1&)8 ΙΠΓΣ ΘΚ 50 ΙΙΥΙ≅ΙΙΥΕ8
Hirata, A. and Shimoda, C. ΒDΕΕΣΓ8 W)∃.∗).∃1/ −(2+,+∗1)+(∀ (, &∋+∀2/% ∋(/% =(2+%& 2.∃+∀3
−%+(&+& !! +& %&&%∀)+1/ ,(∃ )4% ∀(∃−1/ ,(∃−1)+(∀ (, 1&∗(&∋(∃%& +∀ W∗4+;(&1∗∗41∃(−:∗%&
∋(−=%β ./)∃1&)∃.∗).∃1/ 1∀1/:&+& (, &∋( −.)1∀)&8 Οe&st 100 DΥΙ≅DΦΙ8
Hodges, A. R., [rementsova, E. B. and Trybus, [. M. ΒΜΝΝΦΓ8 W4%Ι∋ =+∀2& )( )4% ∃(2
(, :%1&) −:(&+∀ α 1∀2 ∋∃%7%∀)& +) ,∃(− 2+−%∃+;+∀30 ,(∃−+∀3 1 &+∀3/%≅4%12%2 −()(∃
∗(−∋/%68 ΘΚ Γio%Κ Χ/e∗Κ 2830 _ΕΝ_≅_ΕDΣ8
Ikemoto, S., Nakamura, T., [ubo, M. and Shimoda, C. ΒΜΝΝΝΓ8 W8 ∋(−=% &∋(∃./1)+(∀≅
&∋%∗+,+∗ ∗(+/%2≅∗(+/ ∋∃()%+∀ W∋(DΚ∋ +& /(∗1/+;%2 )( )4% &∋+∀2/% ∋(/% =(2: 1∀2 %&&%∀)+1/
,(∃ +)& −(2+,+∗1)+(∀8 ΘΚ Χe%% −ciΚ 1130 ΚΣΚ≅ΚΚΣ8
Itadani, A., Nakamura, T. and Shimoda, C. ΒΜΝΝ_Γ8 φ(∗1/+;1)+(∀ (, ):∋% ! −:(&+∀ 1∀2
Α≅1∗)+∀ )( )4% /%12+∀3 %23% ∃%3+(∀ (, )4% ,(∃%&∋(∃% −%−=∃1∀% +∀ W∗4+;(&1∗∗41∃(−:∗%&
∋(−=%8 Χe%% −tructΚ ΛunctΚ 310 DΦD≅DΕΚ8
Izawa, D., Goto, M., Ξamashita, A., Ξamano, H. and Ξamamoto, M. ΒΜΝΝΚΓ8 Α+&&+(∀
:%1&) Ξ%&D∋ %∀&.∃%& )4% (∀&%) (, −%+(&+& !! =: =/(∗>+∀3 2%3∃121)+(∀ (, ∗:∗/+∀ Ρ2∗DΙ∋8
Μ&ture 4340 ΚΜΕ≅ΚΙΙ8
Jin, ⊥. Ο., Pidoux, A. L., Decker, C., Allshire, R. C. and Fleig, U. ΒΜΝΝΜΓ8 94% −1/Μ∋
∋∃()%+∀ +& 1∀ %&&%∀)+1/ ∗(−∋(∀%∀) (, )4% ,+&&+(∀ :%1&) ∗%∀)∃(−%∃%8 Πo%Κ Χe%%Κ Γio%Κ 220
ΥD_Φ≅ΥDΦΙ8
[jaerulff, S., Lautrup-Larsen, I., Truelsen, S., Pedersen, M. and Nielsen, O. ΒΜΝΝΚΓ8
Ρ(∀&)+).)+7% 1∗)+71)+(∀ (, )4% ,+&&+(∀ :%1&) ∋4%∃(−(∀%≅∃%&∋(∀&+7% ∋1)451: +∀2.∗%& %∗)(∋+∗
−%+(&+& 1∀2 ∃%7%1/& &)%DD 1& 1 −+)(3%∀≅1∗)+71)%2 ∋∃()%+∀ >+∀1&% )1∃3%)8 Πo%Κ Χe%%Κ Γio%Κ
250 ΜΝΣΚ≅ΜΝΚΕ8
[rapp, A., Schmidt, S., Cano, E. and Simanis, &. ΒΜΝΝDΓ8 W8 ∋(−=% ∗2∗DD∋0 )(3%)4%∃
5+)4 &+2Σ∋0 ∋∃(7+2%& 1∀ 1∀∗4(∃ ,(∃ &%∋)1)+(∀ +∀+)+1)+(∀ ∀%)5(∃> ∋∃()%+∀& (∀ )4% &∋+∀2/%
∋(/% =(2:8 ΧurrΚ Γio%Κ 110 DΚΚΕ≅DΚ_Φ8
[rapp, A., Collin, P., Cokoja, A., Dischinger, S., Cano, E. and Simanis, &. ΒΜΝΝ_Γ8 94%
W∗4+;(&1∗∗41∃(−:∗%& ∋(−=% &%∋)1)+(∀ +∀+)+1)+(∀ ∀%)5(∃> ΒW!ΨΓ +& ∃%<.+∃%2 ,(∃ &∋(∃%
,(∃−1)+(∀ +∀ −%+(&+&8 ΘΚ Χe%% −ciΚ 11χ0 ΜΦΦΜ≅ΜΦΕD8
[rementsova, E. B., Hodges, A. R., Lu, H. and Trybus, [. M. ΒΜΝΝ_Γ8 Χ∃(∗%&&+7+): (,
∗4+−%∃+∗ ∗/1&& α −:(&+∀&8 ΘΚ Γio%Κ Χ/e∗Κ 2810 _ΝΥΕ≅_ΝΦ_8
[urahashi, H., Imai, Ξ. and Ξamamoto, M. ΒΜΝΝΜΓ8 9∃(∋(−:(&+∀ +& ∃%<.+∃%2 ,(∃ )4% ∗%//
,.&+(∀ ∋∃(∗%&& 2.∃+∀3 ∗(∀?.31)+(∀ +∀ ,+&&+(∀ :%1&)8 >enes Χe%%s Σ0 ΙΥΚ≅ΙΦΣ8
Martin-Cuadrado, A. B., Duenas, E., Sipiczki, M., &azquez de Aldana, C. R. and del
Rey, F. ΒΜΝΝΙΓ8 94% %∀2(≅=%)1≅D0Ι≅3/.∗1∀1&% %∀3D∋ +& ∃%<.+∃%2 ,(∃ 2+&&(/.)+(∀ (, )4%
∋∃+−1∃: &%∋).− 2.∃+∀3 ∗%// &%∋1∃1)+(∀ +∀ W∗4+;(&1∗∗41∃(−:∗%& ∋(−=%8 ΘΚ Χe%% −ciΚ 11Ρ0
D_ΦΕ≅D_ΕΦ8
Martin-Cuadrado, A. B., Morrell, J. L., [onomi, M., An, H., Petit, C., Osumi, M.,
Balasubramanian, M., Gould, [. L., Del Rey, F. and de Aldana, C. R. ΒΜΝΝΚΓ8 Τ(/%
(, &%∋)+∀& 1∀2 )4% %6(∗:&) ∗(−∋/%6 +∀ )4% ,.∀∗)+(∀ (, 4:2∃(/:)+∗ %∀;:−%& ∃%&∋(∀&+=/%
,(∃ ,+&&+(∀ :%1&) ∗%// &%∋1∃1)+(∀8 Πo%Κ Γio%Κ Χe%% 1Ρ0 ΣΦ_Υ≅ΣΦΦD8
Mata, J., Lyne, R., Burns, G. and Bahler, J. ΒΜΝΝΜΓ8 94% )∃1∀&∗∃+∋)+(∀1/ ∋∃(3∃1− (, −%+(&+&
1∀2 &∋(∃./1)+(∀ +∀ ,+&&+(∀ :%1&)8 Μ&tΚ >enetΚ 320 DΣΙ≅DΣΥ8
Mehta, A. D., Rock, R. S., Rief, M., Spudich, J. A., Mooseker, M. S. and Cheney, R.
E. ΒDΕΕΕΓ8 Ξ:(&+∀≅α +& 1 ∋∃(∗%&&+7% 1∗)+∀≅=1&%2 −()(∃8 Μ&ture 4000 ΚΕΝ≅ΚΕΙ8
Mermall, &., Post, P. L. and Mooseker, M. S. ΒDΕΕΦΓ8 Ο∀∗(∀7%∀)+(∀1/ −:(&+∀& +∀ ∗%//
−(7%−%∀)0 −%−=∃1∀% )∃1,,+∗0 1∀2 &+3∀1/ )∃1∀&2.∗)+(∀8 −cience 2Σχ0 ΚΜΥ≅ΚΙΙ8
Miki, F., Okazaki, [., Shimanuki, M., Ξamamoto, A., Hiraoka, Ξ. and Niwa, O. ΒΜΝΝΜΓ8
94% DΣ≅>δ1 2:∀%+∀ /+34) ∗41+∀≅,1−+/: ∋∃()%+∀ δ/∗D +& ∃%<.+∃%2 ,(∃ ∃%3./1∃ (&∗+//1)(∃:
∀.∗/%1∃ −(7%−%∀) 1∀2 %,,+∗+%∀) ∃%∗(−=+∀1)+(∀ 2.∃+∀3 −%+()+∗ ∋∃(∋41&% +∀ ,+&&+(∀ :%1&)8
Πo%Κ Γio%Κ Χe%% 130 ΕΙΝ≅ΕΣ_8
Molero, G., Cid, &. J., &ivar, C., Nombela, C. and Sanchez-Perez, M. ΒDΕΕΕΓ8 Ρ1∀2+21
1/=+∗1∀& %6(3/.∗1∀1&% 1& 1 ∃%∋(∃)%∃ 3%∀% +∀ W∗4+;(&1∗∗41∃(−:∗%& ∋(−=%8 ΛΙΠ−
Πicro4io%Κ ΤettΚ 1Σ50 DΣΙ≅DΣΦ8
Moreno, S., [lar, A. and Nurse, P. ΒDΕΕDΓ8 Ξ(/%∗./1∃ 3%∀%)+∗ 1∀1/:&+& (, ,+&&+(∀ :%1&)
W∗4+;(&1∗∗41∃(−:∗%& ∋(−=%8 Πet/o,s Ιn0y∗o%Κ 1χ40 ΥΕΚ≅ΦΜΙ8
Niccoli, T., Ξamashita, A., Nurse, P. and Ξamamoto, M. ΒΜΝΝΣΓ8 94% ∋DΚΝ≅ς/.%2 W&−Σ∋
∃%3./1)%& −+∗∃().=./1∃ 2:∀1−+∗& 1∀2 ∀.∗/%1∃ −(7%−%∀) +∀ ,+&&+(∀ :%1&)8 ΘΚ Χe%% −ciΚ
11Σ0 ΚΚΣΙ≅ΚΚΚ_8
Ozoe, F., [urokawa, R., [obayashi, Ξ., Jeong, H. T., Tanaka, [., Sen, [., Nakagawa,
T., Matsuda, H. and [awamukai, M. ΒΜΝΝΜΓ8 94% DΣ≅Ι≅Ι ∋∃()%+∀& Τ12ΜΣ 1∀2 Τ12ΜΚ
∀%31)+7%/: ∃%3./1)% ε:∃Μ =: 1,,%∗)+∀3 +)& /(∗1/+;1)+(∀ +∀ W∗4+;(&1∗∗41∃(−:∗%& ∋(−=%8
Πo%Κ Χe%%Κ Γio%Κ 220 ΥDΝΚ≅ΥDDΕ8
Petersen, J., Οeilguny, D., Egel, R. and Nielsen, O. ΒDΕΕΚΓ8 Ρ41∃1∗)%∃+;1)+(∀ (, ,.&D (,
W∗4+;(&1∗∗41∃(−:∗%& ∋(−=%β 1 2%7%/(∋−%∀)1//: ∗(∀)∃(//%2 ,.∀∗)+(∀ ∀%%2%2 ,(∃
∗(∀?.31)+(∀8 Πo%Κ Χe%%Κ Γio%Κ 150 Ι_ΕΥ≅ΙΥΝΥ8
Petersen, J., Heitz, M. J. and Hagan, I. M. ΒDΕΕΦ1Γ8 Ρ(∀?.31)+(∀ +∀ W8 ∋(−=%β
+2%∀)+,+∗1)+(∀ (, 1 −+∗∃().=./%≅(∃31∀+&+∀3 ∗%∀)∃%0 1 ∃%<.+∃%−%∀) ,(∃ −+∗∃().=./%& 1∀2 1
∃(/% ,(∃ Ξ12Μ8 ΧurrΚ Γio%Κ 80 Ε_Ι≅Ε__8
Petersen, J., Nielsen, O., Egel, R. and Hagan, I. M. ΒDΕΕΦ=Γ8 Α≅1∗)+∀ 2+&)∃+=.)+(∀ 1∀2
,.∀∗)+(∀ 2.∃+∀3 &%6.1/ 2+,,%∃%∀)+1)+(∀ +∀ W∗4+;(&1∗∗41∃(−:∗%& ∋(−=%8 ΘΚ Χe%% −ciΚ 1110
Φ_Υ≅ΦΥ_8
Petersen, J., Nielsen, O., Egel, R. and Hagan, I. M. ΒDΕΕΦ∗Γ8 ΑΗΙ0 1 2(−1+∀ ,(.∀2 +∀
,(∃−+∀&0 )1∃3%)& )4% ,+&&+(∀ :%1&) ,(∃−+∀ Α.&D )( )4% ∋∃(?%∗)+(∀ )+∋ 2.∃+∀3 ∗(∀?.31)+(∀8
ΘΚ Χe%% Γio%Κ 1410 DΜDΥ≅DΜΜΦ8
Pruyne, D. Ο., Schott, D. H. and Bretscher, A. ΒDΕΕΦΓ8 9∃(∋(−:(&+∀≅∗(∀)1+∀+∀3 1∗)+∀
∗1=/%& 2+∃%∗) )4% Ξ:(Μ∋≅2%∋%∀2%∀) ∋(/1∃+;%2 2%/+7%∃: (, &%∗∃%)(∃: 7%&+∗/%& +∀ =.22+∀3
:%1&)8 ΘΚ Χe%% Γio%Κ 1430 DΕΙD≅DΕΣΚ8
Reck-Peterson, S. L., Novick, P. J. and Mooseker, M. S. ΒDΕΕΕΓ8 94% )1+/ (, 1 :%1&) ∗/1&&
α −:(&+∀0 −:(Μ∋0 ,.∀∗)+(∀& 1& 1 /(∗1/+;1)+(∀ 2(−1+∀8 Πo%Κ Γio%Κ Χe%% 100 DΝΝD≅DΝDΥ8
Reck-Peterson, S. L., Tyska, M. J., Novick, P. J. and Mooseker, M. S. ΒΜΝΝDΓ8 94% :%1&)
∗/1&& α −:(&+∀&0 Ξ:(Μ∋ 1∀2 Ξ:(Σ∋0 1∃% ∀(∀∋∃(∗%&&+7% 1∗)+∀≅=1&%2 −()(∃&8 ΘΚ Χe%% Γio%Κ
1530 DDΜD≅DDΜ_8
4340
Journal of Cell Science 122 (23)
Journal of Cell Science
Robinow, C. F. ΒDΕΥΥΓ8 94% ∀.−=%∃ (, ∗4∃(−(&(−%& +∀ WΡΗ!γ∴WΡΡΗΤ∴ΞΖΡ]W
Χ∴Ξε]β /+34) −+∗∃(&∗(∋: (, &)1+∀%2 ∋∃%∋1∃1)+(∀&8 >enetics 8Σ0 ΣΕD≅ΣΕΥ8
Rustici, G., Mata, J., [ivinen, [., Lio, P., Penkett, C. J., Burns, G., Hayles, J., Brazma,
A., Nurse, P. and Bahler, J. ΒΜΝΝΣΓ8 Χ%∃+(2+∗ 3%∀% %6∋∃%&&+(∀ ∋∃(3∃1− (, )4% ,+&&+(∀
:%1&) ∗%// ∗:∗/%8 Μ&tΚ >enetΚ 3Ρ0 ΦΝΕ≅ΦDΥ8
Schott, D. H., Collins, R. N. and Bretscher, A. ΒΜΝΝΜΓ8 W%∗∃%)(∃: 7%&+∗/% )∃1∀&∋(∃) 7%/(∗+):
+∀ /+7+∀3 ∗%//& 2%∋%∀2& (∀ )4% −:(&+∀≅α /%7%∃ 1∃− /%∀3)48 ΘΚ Χe%% Γio%Κ 15Ρ0 ΙΚ≅ΙΕ8
Schuh, M. and Ellenberg, J. ΒΜΝΝΦΓ8  ∀%5 −(2%/ ,(∃ 1&:−−%)∃+∗ &∋+∀2/% ∋(&+)+(∀+∀3 +∀
−(.&% ((∗:)%&8 ΧurrΚ Γio%Κ 180 DΕΦ_≅DΕΕΜ8
Shaner, N. C., Campbell, R. E., Steinbach, P. A., Giepmans, B. N., Palmer, A. E. and
Tsien, R. Ξ. ΒΜΝΝΣΓ8 !−∋∃(7%2 −(∀(−%∃+∗ ∃%20 (∃1∀3% 1∀2 :%//(5 ,/.(∃%&∗%∀) ∋∃()%+∀&
2%∃+7%2 ,∃(− δ+&∗(&(−1 &∋8 ∃%2 ,/.(∃%&∗%∀) ∋∃()%+∀8 Μ&tΚ Γiotec/no%Κ 220 DΚ_Υ≅DΚΥΜ8
Shimoda, C. ΒΜΝΝΣΓ8 Α(∃%&∋(∃% −%−=∃1∀% 1&&%−=/: +∀ :%1&)β ∗((∃2+∀1)+∀3 WΧε& 1∀2
−%−=∃1∀% )∃1,,+∗>+∀38 ΘΚ Χe%% −ciΚ 11Σ0 ΙΦΕ≅ΙΕ_8
Shimoda, C., Hirata, A., [ishida, M., Hashida, T. and Tanaka, [. ΒDΕΦΚΓ8
Ρ41∃1∗)%∃+;1)+(∀ (, −%+(&+&≅2%,+∗+%∀) −.)1∀)& =: %/%∗)∃(∀ −+∗∃(&∗(∋: 1∀2 −1∋∋+∀3 (,
,(.∃ %&&%∀)+1/ 3%∀%& +∀ )4% ,+&&+(∀ :%1&) W∗4+;(&1∗∗41∃(−:∗%& ∋(−=%8 Πo%Κ >enΚ >enetΚ
2000 ΜΚΜ≅ΜΚΥ8
Tanaka, [. and Hirata, A. ΒDΕΦΜΓ8 &∗(&∋(∃% 2%7%/(∋−%∀) +∀ )4% ,+&&+(∀ :%1&)&
W∗4+;(&1∗∗41∃(−:∗%& ∋(−=% 1∀2 W8 ?1∋(∀+∗.&8 ΘΚ Χe%% −ciΚ 5Ρ0 Μ_Ι≅ΜΥΕ8
Tanaka, [., [ohda, T., Ξamashita, A., Nonaka, N. and Ξamamoto, M. ΒΜΝΝΚΓ8
Η∃&D∋χΞ∗∋_∋ (∀ )4% −%+()+∗ WΧε (∃31∀+;%& 1&)∃1/ −+∗∃().=./% 1∃∃1:& ,(∃ (&∗+//1)(∃:
∀.∗/%1∃ −(7%−%∀)8 ΧurrΚ Γio%Κ 150 DΣΥΕ≅DΣΦ_8
Tanaka, Ξ., Okuzaki, D., Ξabuta, N., Ξoneki, T. and Nojima, H. ΒΜΝΝΝΓ8 Τ12ΜΣ +& %&&%∀)+1/
,(∃ ∋∃(/+,%∃1)+(∀ (, 2+∋/(+2 ∗%//& +∀ ,+&&+(∀ :%1&)8 ΛΙΓ− ΤettΚ 4Σ20 ΜΚΣ≅ΜΚΦ8
Taxis, C., Maeder, C., Reber, S., Rathfelder, N., Miura, [., Greger, [., Stelzer, E. H.
and [nop, M. ΒΜΝΝ_Γ8 δ:∀1−+∗ (∃31∀+;1)+(∀ (, )4% 1∗)+∀ ∗:)(&>%/%)(∀ 2.∃+∀3 −%+(&+&
1∀2 &∋(∃% ,(∃−1)+(∀ +∀ =.22+∀3 :%1&)8 ?r&99ic Σ0 D_ΜΦ≅D_ΣΜ8
Toth, J., [ovacs, M., Οang, F., Nyitray, L. and Sellers, J. R. ΒΜΝΝΚΓ8 Ξ:(&+∀ α ,∃(−
δ∃(&(∋4+/1 ∃%7%1/& 2+7%∃&+): (, −()(∃ −%∗41∀+&−& 5+)4+∀ )4% −:(&+∀ α ,1−+/:8 ΘΚ Γio%Κ
Χ/e∗Κ 2800 ΙΝΚΕΣ≅ΙΝ_ΝΙ8
Toya, M., Motegi, F., Nakano, [., Mabuchi, I. and Ξamamoto, M. ΒΜΝΝDΓ8 !2%∀)+,+∗1)+(∀
1∀2 ,.∀∗)+(∀1/ 1∀1/:&+& (, )4% 3%∀% ,(∃ ):∋% ! −:(&+∀ +∀ ,+&&+(∀ :%1&)8 >enes Χe%%s Ρ0
DΦΥ≅DΕΕ8
Tsakraklides, &., [rogh, [., Οang, L., Bizario, J. C., Larson, R. E., Espreafico, E. M.
and Οolenski, J. S. ΒDΕΕΕΓ8 W.=∗%//./1∃ /(∗1/+;1)+(∀ (, ςΑΧ≅−:(&+∀≅α +∀ /+7% −(.&%
−%/1∀(∗:)%&8 ΘΚ Χe%% −ciΚ 1120 ΜΦΚΙ≅ΜΦ_Κ8
Οang, H., Tang, δ., Liu, J., Trautmann, S., Balasundaram, D., McCollum, D. and
Balasubramanian, M. [. ΒΜΝΝΜΓ8 94% −./)+∋∃()%+∀ %6(∗:&) ∗(−∋/%6 +& %&&%∀)+1/ ,(∃
∗%// &%∋1∃1)+(∀ +∀ W∗4+;(&1∗∗41∃(−:∗%& ∋(−=%8 Πo%Κ Γio%Κ Χe%% 130 ΚDΚ≅ΚΜΕ8
Οang, Ξ., δu, H. P., Riggs, M., Rodgers, L. and Οigler, M. ΒDΕΕDΓ8 =:∃Μ0 1
W∗4+;(&1∗∗41∃(−:∗%& ∋(−=% 3%∀% %∀∗(2+∀3 1 ∋∃()%+∀ >+∀1&% ∗1∋1=/% (, ∋1∃)+1/
&.∋∋∃%&&+(∀ (, )4% ∃1&D −.)1∀) ∋4%∀():∋%8 Πo%Κ Χe%%Κ Γio%Κ 110 ΙΚΚΣ≅ΙΚ_Ι8
Οeber, I., Gruber, C. and Steinberg, G. ΒΜΝΝΙΓ8  ∗/1&&≅α −:(&+∀ ∃%<.+∃%2 ,(∃ −1)+∀30
4:∋41/ 3∃(5)40 1∀2 ∋1)4(3%∀+∗+): +∀ )4% 2+−(∃∋4+∗ ∋/1∀) ∋1)4(3%∀ Ο&)+/13( −1:2+&8 ϑ%&nt
Χe%% 150 ΜΦΜ_≅ΜΦΣΜ8
Οeber, [. L., Sokac, A. M., Berg, J. S., Cheney, R. E. and Bement, Ο. M. ΒΜΝΝΣΓ8 
−+∗∃().=./%≅=+∀2+∀3 −:(&+∀ ∃%<.+∃%2 ,(∃ ∀.∗/%1∃ 1∀∗4(∃+∀3 1∀2 &∋+∀2/% 1&&%−=/:8 Μ&ture
4310 ΙΜΚ≅ΙΜΕ8
Οin, T. Z., Gachet, Ξ., Mulvihill, D. P., May, [. M. and Hyams, J. S. ΒΜΝΝDΓ8 95( ):∋%
α −:(&+∀& 5+)4 ∀(∀≅(7%∃/1∋∋+∀3 ,.∀∗)+(∀& +∀ )4% ,+&&+(∀ :%1&) W∗4+;(&1∗∗41∃(−:∗%&
∋(−=%β Ξ:(ΚΜ +& ∗(∀∗%∃∀%2 5+)4 3∃(5)4 ∋(/1∃+): 1∀2 ∗:)(>+∀%&+&0 Ξ:(ΚD +& 1 ∗(−∋(∀%∀)
(, )4% ∗:)(>+∀%)+∗ 1∗)+∀ ∃+∀38 ΘΚ Χe%% −ciΚ 1140 _Ε≅ΥΕ8
Ξamamoto, A., Οest, R. R., McIntosh, J. R. and Hiraoka, Ξ. ΒDΕΕΕΓ8  ∗:)(∋/1&−+∗
2:∀%+∀ 4%17: ∗41+∀ +& ∃%<.+∃%2 ,(∃ (&∗+//1)(∃: ∀.∗/%1∃ −(7%−%∀) (, −%+()+∗ ∋∃(∋41&%
1∀2 %,,+∗+%∀) −%+()+∗ ∃%∗(−=+∀1)+(∀ +∀ ,+&&+(∀ :%1&)8 ΘΚ Χe%% Γio%Κ 1450 DΜΙΙ≅DΜΣΕ8