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![Page 1: Model-Convolution Approach to Modeling Green Fluorescent Protein Dynamics: Application to Yeast Cell Division David Odde Dept. of Biomedical Engineering.](https://reader030.fdocuments.in/reader030/viewer/2022032517/56649cac5503460f9496ed95/html5/thumbnails/1.jpg)
Model-Convolution Approach to Modeling Green Fluorescent
Protein Dynamics: Application to Yeast Cell Division
David Odde
Dept. of Biomedical Engineering
University of Minnesota
![Page 2: Model-Convolution Approach to Modeling Green Fluorescent Protein Dynamics: Application to Yeast Cell Division David Odde Dept. of Biomedical Engineering.](https://reader030.fdocuments.in/reader030/viewer/2022032517/56649cac5503460f9496ed95/html5/thumbnails/2.jpg)
Mitotic Spindle
spindle pole
chromosomes
kinetochore
1.7 µmIn budding yeast:
~40 MTs10-20 µm
In animal cells:
~1000 MTs
interpolarmicrotubule
- -
++
++
kinetochore microtubule
bifunctionalplus-end motors
+ +
spindle pole
COMPRESSION
TENSION
![Page 3: Model-Convolution Approach to Modeling Green Fluorescent Protein Dynamics: Application to Yeast Cell Division David Odde Dept. of Biomedical Engineering.](https://reader030.fdocuments.in/reader030/viewer/2022032517/56649cac5503460f9496ed95/html5/thumbnails/3.jpg)
Microtubule Dynamic Instability
![Page 4: Model-Convolution Approach to Modeling Green Fluorescent Protein Dynamics: Application to Yeast Cell Division David Odde Dept. of Biomedical Engineering.](https://reader030.fdocuments.in/reader030/viewer/2022032517/56649cac5503460f9496ed95/html5/thumbnails/4.jpg)
Leng
th (
µm
)
Time (minutes)
“Catastrophe”
“Rescue”
Microtubule “Dynamic Instability”
Vg
Vs
kc
kr
Hypothesis: The kinetochore modulates the DI parameters
![Page 5: Model-Convolution Approach to Modeling Green Fluorescent Protein Dynamics: Application to Yeast Cell Division David Odde Dept. of Biomedical Engineering.](https://reader030.fdocuments.in/reader030/viewer/2022032517/56649cac5503460f9496ed95/html5/thumbnails/5.jpg)
Can only get peaks here
Not here
MT Length Distribution for Pure Dynamic Instability
Right PoleLeft Pole
1.7
![Page 6: Model-Convolution Approach to Modeling Green Fluorescent Protein Dynamics: Application to Yeast Cell Division David Odde Dept. of Biomedical Engineering.](https://reader030.fdocuments.in/reader030/viewer/2022032517/56649cac5503460f9496ed95/html5/thumbnails/6.jpg)
Budding Yeast Spindle Geometry
![Page 7: Model-Convolution Approach to Modeling Green Fluorescent Protein Dynamics: Application to Yeast Cell Division David Odde Dept. of Biomedical Engineering.](https://reader030.fdocuments.in/reader030/viewer/2022032517/56649cac5503460f9496ed95/html5/thumbnails/7.jpg)
Congression in S. cerevisiae
P PEQ
Green=Cse4-GFP kMT Plus Ends
Red=Spc29-CFP kMT Minus Ends
![Page 8: Model-Convolution Approach to Modeling Green Fluorescent Protein Dynamics: Application to Yeast Cell Division David Odde Dept. of Biomedical Engineering.](https://reader030.fdocuments.in/reader030/viewer/2022032517/56649cac5503460f9496ed95/html5/thumbnails/8.jpg)
“Experiment-Deconvolution”vs. “Model-Convolution”
Model Experiment
Deconvolution
Convolution
![Page 9: Model-Convolution Approach to Modeling Green Fluorescent Protein Dynamics: Application to Yeast Cell Division David Odde Dept. of Biomedical Engineering.](https://reader030.fdocuments.in/reader030/viewer/2022032517/56649cac5503460f9496ed95/html5/thumbnails/9.jpg)
Point Spread Function (PSF)
• A point source of light is spread via diffraction through a circular aperture
• Modeling needs to account for PSF
-0.4-0.20+0.2+0.4 μm
![Page 10: Model-Convolution Approach to Modeling Green Fluorescent Protein Dynamics: Application to Yeast Cell Division David Odde Dept. of Biomedical Engineering.](https://reader030.fdocuments.in/reader030/viewer/2022032517/56649cac5503460f9496ed95/html5/thumbnails/10.jpg)
Simulated Image Obtainedby Model-Convolution of
Original Distribution
Original FluorophoreDistribution
Image Obtained by Deconvolution
of Simulated Image
Potential Pitfalls of Deconvolution
![Page 11: Model-Convolution Approach to Modeling Green Fluorescent Protein Dynamics: Application to Yeast Cell Division David Odde Dept. of Biomedical Engineering.](https://reader030.fdocuments.in/reader030/viewer/2022032517/56649cac5503460f9496ed95/html5/thumbnails/11.jpg)
Cse4-GFP Fluorescence Distribution
Experimentally Observed
Theoretically Predicted
![Page 12: Model-Convolution Approach to Modeling Green Fluorescent Protein Dynamics: Application to Yeast Cell Division David Odde Dept. of Biomedical Engineering.](https://reader030.fdocuments.in/reader030/viewer/2022032517/56649cac5503460f9496ed95/html5/thumbnails/12.jpg)
Dynamic Instability Only Model
Sprague et al., Biophysical J., 2003
![Page 13: Model-Convolution Approach to Modeling Green Fluorescent Protein Dynamics: Application to Yeast Cell Division David Odde Dept. of Biomedical Engineering.](https://reader030.fdocuments.in/reader030/viewer/2022032517/56649cac5503460f9496ed95/html5/thumbnails/13.jpg)
Modeling ApproachModel
Probability that themodel is consistent with the data
ParameterSpace
(a1, a2, a3,…aN)
<Cutoff?
Experimental Data yes
no
Accept Model
ParameterSpace
Reject Model
ParameterSpace
Accept Model
ParameterSpace
![Page 14: Model-Convolution Approach to Modeling Green Fluorescent Protein Dynamics: Application to Yeast Cell Division David Odde Dept. of Biomedical Engineering.](https://reader030.fdocuments.in/reader030/viewer/2022032517/56649cac5503460f9496ed95/html5/thumbnails/14.jpg)
Modeling ApproachModel assumptions:1) Metaphase kinetochore microtubule dynamics
are at steady-state (not time-dependent)2) One microtubule per kinetochore3) Microtubules never detach from kinetochores4) Parameters can be:• Constant• Spatially-dependent (relative to poles)• Spatially-dependent (relative to sister
kinetochore)
![Page 15: Model-Convolution Approach to Modeling Green Fluorescent Protein Dynamics: Application to Yeast Cell Division David Odde Dept. of Biomedical Engineering.](https://reader030.fdocuments.in/reader030/viewer/2022032517/56649cac5503460f9496ed95/html5/thumbnails/15.jpg)
“Microtubule Chemotaxis” in a Chemical Gradient
ImmobileKinase
MobilePhosphatase
A: Phosphorylated ProteinB: Dephosphorylated Protein
k*Surface reaction B-->A
kHomogeneous reaction A-->B
KinetochoreMicrotubules
- +
ImmobileKinase
MT Destabilizer
Position
Concentration
X=0 X=L
![Page 16: Model-Convolution Approach to Modeling Green Fluorescent Protein Dynamics: Application to Yeast Cell Division David Odde Dept. of Biomedical Engineering.](https://reader030.fdocuments.in/reader030/viewer/2022032517/56649cac5503460f9496ed95/html5/thumbnails/16.jpg)
Could tension stabilize kinetochore microtubules?
Tension
Kip3
![Page 17: Model-Convolution Approach to Modeling Green Fluorescent Protein Dynamics: Application to Yeast Cell Division David Odde Dept. of Biomedical Engineering.](https://reader030.fdocuments.in/reader030/viewer/2022032517/56649cac5503460f9496ed95/html5/thumbnails/17.jpg)
Distribution of Cse4-GFP: Catastophe Gradient with Tension Between Sister Kinetochore-Dependent Rescue
![Page 18: Model-Convolution Approach to Modeling Green Fluorescent Protein Dynamics: Application to Yeast Cell Division David Odde Dept. of Biomedical Engineering.](https://reader030.fdocuments.in/reader030/viewer/2022032517/56649cac5503460f9496ed95/html5/thumbnails/18.jpg)
Model Combinations
![Page 19: Model-Convolution Approach to Modeling Green Fluorescent Protein Dynamics: Application to Yeast Cell Division David Odde Dept. of Biomedical Engineering.](https://reader030.fdocuments.in/reader030/viewer/2022032517/56649cac5503460f9496ed95/html5/thumbnails/19.jpg)
123
Catastrophe Gradient-Tension Rescue Model
![Page 20: Model-Convolution Approach to Modeling Green Fluorescent Protein Dynamics: Application to Yeast Cell Division David Odde Dept. of Biomedical Engineering.](https://reader030.fdocuments.in/reader030/viewer/2022032517/56649cac5503460f9496ed95/html5/thumbnails/20.jpg)
Conclusions
• Congression in budding yeast is mediated by:– Spatially-dependent catastrophe
gradient– Tension between sister kinetochore-
dependent rescue
• Model-convolution can be a useful tool for comparing fluorescent microscopy data to model predictions
![Page 21: Model-Convolution Approach to Modeling Green Fluorescent Protein Dynamics: Application to Yeast Cell Division David Odde Dept. of Biomedical Engineering.](https://reader030.fdocuments.in/reader030/viewer/2022032517/56649cac5503460f9496ed95/html5/thumbnails/21.jpg)
Acknowledgements
• Melissa Gardner, Brian Sprague (Uof M)
• Chad Pearson, Paul Maddox, Kerry Bloom,Ted Salmon (UNC-CH)
• National Science Foundation
• Whitaker Foundation
• McKnight Foundation
![Page 22: Model-Convolution Approach to Modeling Green Fluorescent Protein Dynamics: Application to Yeast Cell Division David Odde Dept. of Biomedical Engineering.](https://reader030.fdocuments.in/reader030/viewer/2022032517/56649cac5503460f9496ed95/html5/thumbnails/22.jpg)
Simulated Image Obtainedby Convolution of PSF and GWN
with Original Distribution
Original FluorophoreDistribution
Model-Convolution
![Page 23: Model-Convolution Approach to Modeling Green Fluorescent Protein Dynamics: Application to Yeast Cell Division David Odde Dept. of Biomedical Engineering.](https://reader030.fdocuments.in/reader030/viewer/2022032517/56649cac5503460f9496ed95/html5/thumbnails/23.jpg)
Kinetochore MT Lengths in Budding Yeast
Experimentally Observed
Theoretically Predicted
?
2 µm
![Page 24: Model-Convolution Approach to Modeling Green Fluorescent Protein Dynamics: Application to Yeast Cell Division David Odde Dept. of Biomedical Engineering.](https://reader030.fdocuments.in/reader030/viewer/2022032517/56649cac5503460f9496ed95/html5/thumbnails/24.jpg)
Catastrophe Gradient Model
Fre
quen
cy (
min
-1)
Normalized Spindle Position
Sprague et al., Biophys. J., 2003
![Page 25: Model-Convolution Approach to Modeling Green Fluorescent Protein Dynamics: Application to Yeast Cell Division David Odde Dept. of Biomedical Engineering.](https://reader030.fdocuments.in/reader030/viewer/2022032517/56649cac5503460f9496ed95/html5/thumbnails/25.jpg)
Distribution of Cse4-GFP: Catastrophe Gradient Model
![Page 26: Model-Convolution Approach to Modeling Green Fluorescent Protein Dynamics: Application to Yeast Cell Division David Odde Dept. of Biomedical Engineering.](https://reader030.fdocuments.in/reader030/viewer/2022032517/56649cac5503460f9496ed95/html5/thumbnails/26.jpg)
Experimental Cse4-GFP FRAP
•Cse4-GFP does not turnover on kinetochore
•Kinetochores rarely persist in opposite half-spindle
Pearson et al., Current Biology, in press
![Page 27: Model-Convolution Approach to Modeling Green Fluorescent Protein Dynamics: Application to Yeast Cell Division David Odde Dept. of Biomedical Engineering.](https://reader030.fdocuments.in/reader030/viewer/2022032517/56649cac5503460f9496ed95/html5/thumbnails/27.jpg)
Cse4-GFP FRAP: Modeling and Experiment
Catastrophe Gradient Simulation
Experiment
![Page 28: Model-Convolution Approach to Modeling Green Fluorescent Protein Dynamics: Application to Yeast Cell Division David Odde Dept. of Biomedical Engineering.](https://reader030.fdocuments.in/reader030/viewer/2022032517/56649cac5503460f9496ed95/html5/thumbnails/28.jpg)
Cse4-GFP FRAP: Modeling and Experiment
![Page 29: Model-Convolution Approach to Modeling Green Fluorescent Protein Dynamics: Application to Yeast Cell Division David Odde Dept. of Biomedical Engineering.](https://reader030.fdocuments.in/reader030/viewer/2022032517/56649cac5503460f9496ed95/html5/thumbnails/29.jpg)
Gradients in Phospho-state
1.0
0.8
0.6
0.4
0.2
0.0
Con
cen
trati
on
, Y
1.00.80.60.40.20.0
Position, X
If k= 50 s-1, D=5 µm2/s, and L=1 µm, then =3
MT Destabilizer
Position
Concentration
X=0 X=L
![Page 30: Model-Convolution Approach to Modeling Green Fluorescent Protein Dynamics: Application to Yeast Cell Division David Odde Dept. of Biomedical Engineering.](https://reader030.fdocuments.in/reader030/viewer/2022032517/56649cac5503460f9496ed95/html5/thumbnails/30.jpg)
Could tension stabilize kinetochore microtubules?
TensionTension
Kip3
![Page 31: Model-Convolution Approach to Modeling Green Fluorescent Protein Dynamics: Application to Yeast Cell Division David Odde Dept. of Biomedical Engineering.](https://reader030.fdocuments.in/reader030/viewer/2022032517/56649cac5503460f9496ed95/html5/thumbnails/31.jpg)
Catastophe Gradient with Tension Between Sister Kinetochore-Dependent Rescue Model
![Page 32: Model-Convolution Approach to Modeling Green Fluorescent Protein Dynamics: Application to Yeast Cell Division David Odde Dept. of Biomedical Engineering.](https://reader030.fdocuments.in/reader030/viewer/2022032517/56649cac5503460f9496ed95/html5/thumbnails/32.jpg)
Experimental Cse4-GFP in Cdc6 mutants
WT Cdc6
![Page 33: Model-Convolution Approach to Modeling Green Fluorescent Protein Dynamics: Application to Yeast Cell Division David Odde Dept. of Biomedical Engineering.](https://reader030.fdocuments.in/reader030/viewer/2022032517/56649cac5503460f9496ed95/html5/thumbnails/33.jpg)
Cse4-GFP in Cdc6 Cells: No tension between sister kinetochores
Rescue Gradient with Tension-Dependent Catastrophe Model (No Tension)
Normalized Spindle Position
Fre
quen
cy (
min
-1)
Catastrophe Gradient with Tension-Dependent Rescue Model (No Tension)
Fre
quen
cy (
min
-1)
Normalized Spindle Position
![Page 34: Model-Convolution Approach to Modeling Green Fluorescent Protein Dynamics: Application to Yeast Cell Division David Odde Dept. of Biomedical Engineering.](https://reader030.fdocuments.in/reader030/viewer/2022032517/56649cac5503460f9496ed95/html5/thumbnails/34.jpg)
Cse4-GFP in Cdc6 Cells: No tension between sister kinetochores
0.022
0.023
0.024
0.025
0.026
0.027
0.028
0 0.1 0.2 0.3 0.4 0.5 0.6 0.7 0.8 0.9 1
Normalized Spindle Position
Frac
tion
Fluo
resc
ence
Experimental cdc6 mutants- No Replication (n=27)Catastrophe Gradient with Tension-Dep. Rescue (No Tension); p=0.11Rescue Gradient with Tension-Dep. Catastrophe (No Tension); p<<.01
![Page 35: Model-Convolution Approach to Modeling Green Fluorescent Protein Dynamics: Application to Yeast Cell Division David Odde Dept. of Biomedical Engineering.](https://reader030.fdocuments.in/reader030/viewer/2022032517/56649cac5503460f9496ed95/html5/thumbnails/35.jpg)
Rescue Gradient Model
Normalized Spindle Position
Ca
tast
rop
he
or
Re
scu
e F
requ
enc
y (m
in-1)
![Page 36: Model-Convolution Approach to Modeling Green Fluorescent Protein Dynamics: Application to Yeast Cell Division David Odde Dept. of Biomedical Engineering.](https://reader030.fdocuments.in/reader030/viewer/2022032517/56649cac5503460f9496ed95/html5/thumbnails/36.jpg)
Simulation of Budding Yeast Mitosis
Metaphase AnaphasePrometaphase
Start with randompositions, let simulationreach steady-state
Eliminate cohesion,set spring constant to 0
![Page 37: Model-Convolution Approach to Modeling Green Fluorescent Protein Dynamics: Application to Yeast Cell Division David Odde Dept. of Biomedical Engineering.](https://reader030.fdocuments.in/reader030/viewer/2022032517/56649cac5503460f9496ed95/html5/thumbnails/37.jpg)
MINIMUM ABSOLUTE SISTER KINETOCHORE SEPARATION DISTANCE
![Page 38: Model-Convolution Approach to Modeling Green Fluorescent Protein Dynamics: Application to Yeast Cell Division David Odde Dept. of Biomedical Engineering.](https://reader030.fdocuments.in/reader030/viewer/2022032517/56649cac5503460f9496ed95/html5/thumbnails/38.jpg)
WT Stu2p-depleted
Pearson et al., Mol. Biol. Cell, 2003
Stu2p-mediated catastrophe gradient?
![Page 39: Model-Convolution Approach to Modeling Green Fluorescent Protein Dynamics: Application to Yeast Cell Division David Odde Dept. of Biomedical Engineering.](https://reader030.fdocuments.in/reader030/viewer/2022032517/56649cac5503460f9496ed95/html5/thumbnails/39.jpg)
Green Fluorescent Protein
![Page 40: Model-Convolution Approach to Modeling Green Fluorescent Protein Dynamics: Application to Yeast Cell Division David Odde Dept. of Biomedical Engineering.](https://reader030.fdocuments.in/reader030/viewer/2022032517/56649cac5503460f9496ed95/html5/thumbnails/40.jpg)
![Page 41: Model-Convolution Approach to Modeling Green Fluorescent Protein Dynamics: Application to Yeast Cell Division David Odde Dept. of Biomedical Engineering.](https://reader030.fdocuments.in/reader030/viewer/2022032517/56649cac5503460f9496ed95/html5/thumbnails/41.jpg)
M
D
Prometaphase Spindles and the Importance of Tension in Mitosis
“Syntely”
Ipl1-mediated detachment of kinetochores under low tension
Dewar et al., Nature 2004
![Page 42: Model-Convolution Approach to Modeling Green Fluorescent Protein Dynamics: Application to Yeast Cell Division David Odde Dept. of Biomedical Engineering.](https://reader030.fdocuments.in/reader030/viewer/2022032517/56649cac5503460f9496ed95/html5/thumbnails/42.jpg)
![Page 43: Model-Convolution Approach to Modeling Green Fluorescent Protein Dynamics: Application to Yeast Cell Division David Odde Dept. of Biomedical Engineering.](https://reader030.fdocuments.in/reader030/viewer/2022032517/56649cac5503460f9496ed95/html5/thumbnails/43.jpg)
![Page 44: Model-Convolution Approach to Modeling Green Fluorescent Protein Dynamics: Application to Yeast Cell Division David Odde Dept. of Biomedical Engineering.](https://reader030.fdocuments.in/reader030/viewer/2022032517/56649cac5503460f9496ed95/html5/thumbnails/44.jpg)
MT Length Distributions•Regard MT dynamic instability as diffusion + drift•The drift velocity is a constant given by
•For constant Vg, Vs, kc, and kr, the length distribution is exponential
p x ~ eVdDx
Vd<0 exponential decayVd>0 exponential growth
Vd x Lg Lstc
Vg tg Vststg ts
Vgkc
Vs kr1kc
1kr
![Page 45: Model-Convolution Approach to Modeling Green Fluorescent Protein Dynamics: Application to Yeast Cell Division David Odde Dept. of Biomedical Engineering.](https://reader030.fdocuments.in/reader030/viewer/2022032517/56649cac5503460f9496ed95/html5/thumbnails/45.jpg)
Sister Kinetochore Microtubule Dynamics
![Page 46: Model-Convolution Approach to Modeling Green Fluorescent Protein Dynamics: Application to Yeast Cell Division David Odde Dept. of Biomedical Engineering.](https://reader030.fdocuments.in/reader030/viewer/2022032517/56649cac5503460f9496ed95/html5/thumbnails/46.jpg)
Simulated Image Obtainedby Convolution of PSF and GWN
with Original Distribution
Original FluorophoreDistribution
Model-Convolution
![Page 47: Model-Convolution Approach to Modeling Green Fluorescent Protein Dynamics: Application to Yeast Cell Division David Odde Dept. of Biomedical Engineering.](https://reader030.fdocuments.in/reader030/viewer/2022032517/56649cac5503460f9496ed95/html5/thumbnails/47.jpg)
“Directional Instability”
Skibbens et al., JCB 1993
![Page 48: Model-Convolution Approach to Modeling Green Fluorescent Protein Dynamics: Application to Yeast Cell Division David Odde Dept. of Biomedical Engineering.](https://reader030.fdocuments.in/reader030/viewer/2022032517/56649cac5503460f9496ed95/html5/thumbnails/48.jpg)
Tension on the kinetochore promotes switching to the growth state?
Skibbens and Salmon, Exp. Cell Res., 1997
![Page 49: Model-Convolution Approach to Modeling Green Fluorescent Protein Dynamics: Application to Yeast Cell Division David Odde Dept. of Biomedical Engineering.](https://reader030.fdocuments.in/reader030/viewer/2022032517/56649cac5503460f9496ed95/html5/thumbnails/49.jpg)
Tension Between Sister Kinetochore-Dependent Rescue
kr kroeF
![Page 50: Model-Convolution Approach to Modeling Green Fluorescent Protein Dynamics: Application to Yeast Cell Division David Odde Dept. of Biomedical Engineering.](https://reader030.fdocuments.in/reader030/viewer/2022032517/56649cac5503460f9496ed95/html5/thumbnails/50.jpg)
Catastrophe Gradient withTension-Rescue Model
Lack of Equator Crossing in the CatastropheGradient with Tension-Rescue Model
~25% FRAP recovery ~5% FRAP recovery
![Page 51: Model-Convolution Approach to Modeling Green Fluorescent Protein Dynamics: Application to Yeast Cell Division David Odde Dept. of Biomedical Engineering.](https://reader030.fdocuments.in/reader030/viewer/2022032517/56649cac5503460f9496ed95/html5/thumbnails/51.jpg)
Microtubule Dynamic Instability
![Page 52: Model-Convolution Approach to Modeling Green Fluorescent Protein Dynamics: Application to Yeast Cell Division David Odde Dept. of Biomedical Engineering.](https://reader030.fdocuments.in/reader030/viewer/2022032517/56649cac5503460f9496ed95/html5/thumbnails/52.jpg)
Model for Chemotactic Gradients of Phosphoprotein State
cAt
D 2cAx2
kcA Fick’s Second Law with First-Order HomogeneousReaction (A->B)
DcAx x0
k *cB 0 B.C. 1: Surface reaction at x=0 (B->A)
DcAx xL
0 B.C. 2: No net flux at x=L
cA cB cT Conservation of phosphoprotein
Sprague et al., Biophys. J., 2003
![Page 53: Model-Convolution Approach to Modeling Green Fluorescent Protein Dynamics: Application to Yeast Cell Division David Odde Dept. of Biomedical Engineering.](https://reader030.fdocuments.in/reader030/viewer/2022032517/56649cac5503460f9496ed95/html5/thumbnails/53.jpg)
Predicted Concentration Profile
where
Y cA cT
X x L
kL2
D
A*e2
e2 1 * 1 e2 B*
e2 1 * 1 e2 * k
*LD
Y Ae X BeX
![Page 54: Model-Convolution Approach to Modeling Green Fluorescent Protein Dynamics: Application to Yeast Cell Division David Odde Dept. of Biomedical Engineering.](https://reader030.fdocuments.in/reader030/viewer/2022032517/56649cac5503460f9496ed95/html5/thumbnails/54.jpg)
Model Predictions: Effect of Surface Reaction Rate
1.0
0.8
0.6
0.4
0.2
0.0
Con
cen
trati
on
, Y
1.00.80.60.40.20.0
Position, X
![Page 55: Model-Convolution Approach to Modeling Green Fluorescent Protein Dynamics: Application to Yeast Cell Division David Odde Dept. of Biomedical Engineering.](https://reader030.fdocuments.in/reader030/viewer/2022032517/56649cac5503460f9496ed95/html5/thumbnails/55.jpg)
Defining “Metaphase” in Budding Yeast