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Motor Max Force Max Speed Max Power R O T A R Y Flagellar motor (8 units, E. coli) 2400 pN nm 95 pN 300 Hz 35 m/s 2000 pN nm @ 150 Hz 1.9 x 10 6 pN nm /s (Vibrio) 1700 Hz 220 m/s (single unit) 300 pN nm 12 pN 300 Hz 35 m/s 250 pN nm @ 150 Hz 2.4 x 10 5 pN nm /s F 1 -ATPase 40 pN nm 40 pN 150 Hz 0.9 m/s 20 pN nm @ 75 Hz 9 x 10 3 pN nm /s L I N E A R Myosin (single molecule in muscle or in vitro) 6 pN 10 m/s (speed of array, each molecule mostly detached) 2 pN @ 10 /s x 20 nm 400 pN nm /s Kinesin 5 pN 1 m/s 2.5 pN @ 0.5 m /s 1.25 x 10 3 pN nm /s

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Page 1: MotorMax ForceMax SpeedMax Power ROTARY ROTARY Flagellar motor (8 units, E. coli) 2400 pN nm 95 pN 300 Hz 35 m/s 2000 pN nm @ 150 Hz 1.9 x 10 6 pN nm

Motor Max Force Max Speed Max Power

ROTARY

Flagellar motor (8 units, E. coli)

2400 pN nm

95 pN

300 Hz

35 m/s

2000 pN nm @ 150 Hz

1.9 x 106 pN nm /s

(Vibrio) 1700 Hz

220 m/s

(single unit) 300 pN nm

12 pN

300 Hz

35 m/s

250 pN nm @ 150 Hz

2.4 x 105 pN nm /s

F1 -ATPase 40 pN nm

40 pN

150 Hz

0.9 m/s

20 pN nm @ 75 Hz

9 x 103 pN nm /s

LINEAR

Myosin (single molecule in muscle or in vitro)

6 pN 10 m/s(speed of array, each

molecule mostly detached)

2 pN @ 10 /s x 20 nm

400 pN nm /s

Kinesin 5 pN 1 m/s 2.5 pN @ 0.5 m /s

1.25 x 103 pN nm /s

RNA polymerase

20 pN 0.01 m/s ~200 pN nm /s

Page 2: MotorMax ForceMax SpeedMax Power ROTARY ROTARY Flagellar motor (8 units, E. coli) 2400 pN nm 95 pN 300 Hz 35 m/s 2000 pN nm @ 150 Hz 1.9 x 10 6 pN nm

Overview of lectures

• BIOLOGY was introduced by Judy Armitage’s

• BIOPHYSICS - Experimental Techniques to measure rotary molecular motors

Flagellar Motor

F1-ATPase

Page 3: MotorMax ForceMax SpeedMax Power ROTARY ROTARY Flagellar motor (8 units, E. coli) 2400 pN nm 95 pN 300 Hz 35 m/s 2000 pN nm @ 150 Hz 1.9 x 10 6 pN nm

Single-molecule experiments on bacterial flagellar motors

Page 4: MotorMax ForceMax SpeedMax Power ROTARY ROTARY Flagellar motor (8 units, E. coli) 2400 pN nm 95 pN 300 Hz 35 m/s 2000 pN nm @ 150 Hz 1.9 x 10 6 pN nm
Page 5: MotorMax ForceMax SpeedMax Power ROTARY ROTARY Flagellar motor (8 units, E. coli) 2400 pN nm 95 pN 300 Hz 35 m/s 2000 pN nm @ 150 Hz 1.9 x 10 6 pN nm

Rotor

Stator

Page 6: MotorMax ForceMax SpeedMax Power ROTARY ROTARY Flagellar motor (8 units, E. coli) 2400 pN nm 95 pN 300 Hz 35 m/s 2000 pN nm @ 150 Hz 1.9 x 10 6 pN nm
Page 7: MotorMax ForceMax SpeedMax Power ROTARY ROTARY Flagellar motor (8 units, E. coli) 2400 pN nm 95 pN 300 Hz 35 m/s 2000 pN nm @ 150 Hz 1.9 x 10 6 pN nm
Page 8: MotorMax ForceMax SpeedMax Power ROTARY ROTARY Flagellar motor (8 units, E. coli) 2400 pN nm 95 pN 300 Hz 35 m/s 2000 pN nm @ 150 Hz 1.9 x 10 6 pN nm

Continuous switch model

Page 9: MotorMax ForceMax SpeedMax Power ROTARY ROTARY Flagellar motor (8 units, E. coli) 2400 pN nm 95 pN 300 Hz 35 m/s 2000 pN nm @ 150 Hz 1.9 x 10 6 pN nm
Page 10: MotorMax ForceMax SpeedMax Power ROTARY ROTARY Flagellar motor (8 units, E. coli) 2400 pN nm 95 pN 300 Hz 35 m/s 2000 pN nm @ 150 Hz 1.9 x 10 6 pN nm

Switching

F Bai, RW. Branch, D Nicolau, TPilizota, BC Steel, PK Maini, RM Berry (2010)Conformational spread as a mechanism for cooperativity in the bacterial flagellar switchScience 327:685-689

Page 11: MotorMax ForceMax SpeedMax Power ROTARY ROTARY Flagellar motor (8 units, E. coli) 2400 pN nm 95 pN 300 Hz 35 m/s 2000 pN nm @ 150 Hz 1.9 x 10 6 pN nm

Bacterial Chemotaxis

Page 12: MotorMax ForceMax SpeedMax Power ROTARY ROTARY Flagellar motor (8 units, E. coli) 2400 pN nm 95 pN 300 Hz 35 m/s 2000 pN nm @ 150 Hz 1.9 x 10 6 pN nm

1-D Ising model of flagellar switch

movie

Page 13: MotorMax ForceMax SpeedMax Power ROTARY ROTARY Flagellar motor (8 units, E. coli) 2400 pN nm 95 pN 300 Hz 35 m/s 2000 pN nm @ 150 Hz 1.9 x 10 6 pN nm

Tethered cells

Cell body rotates at ~ 10 Hz

1 m

Flagellum tethered to coverslip

Cell body

Coverslip in microscope

Page 14: MotorMax ForceMax SpeedMax Power ROTARY ROTARY Flagellar motor (8 units, E. coli) 2400 pN nm 95 pN 300 Hz 35 m/s 2000 pN nm @ 150 Hz 1.9 x 10 6 pN nm

Frequency (Hz)

fast beads, small.mov

Page 15: MotorMax ForceMax SpeedMax Power ROTARY ROTARY Flagellar motor (8 units, E. coli) 2400 pN nm 95 pN 300 Hz 35 m/s 2000 pN nm @ 150 Hz 1.9 x 10 6 pN nm

Work = torque x angle

Torque = d(work) / d(angle)

Low Reynolds number:Torque = viscous drag coefficient x angular velocity

Page 16: MotorMax ForceMax SpeedMax Power ROTARY ROTARY Flagellar motor (8 units, E. coli) 2400 pN nm 95 pN 300 Hz 35 m/s 2000 pN nm @ 150 Hz 1.9 x 10 6 pN nm

Beads attached to the motor

Page 17: MotorMax ForceMax SpeedMax Power ROTARY ROTARY Flagellar motor (8 units, E. coli) 2400 pN nm 95 pN 300 Hz 35 m/s 2000 pN nm @ 150 Hz 1.9 x 10 6 pN nm

Finite, variable switch times

Page 18: MotorMax ForceMax SpeedMax Power ROTARY ROTARY Flagellar motor (8 units, E. coli) 2400 pN nm 95 pN 300 Hz 35 m/s 2000 pN nm @ 150 Hz 1.9 x 10 6 pN nm

Switch times distribution predicted by model

Page 19: MotorMax ForceMax SpeedMax Power ROTARY ROTARY Flagellar motor (8 units, E. coli) 2400 pN nm 95 pN 300 Hz 35 m/s 2000 pN nm @ 150 Hz 1.9 x 10 6 pN nm

… further detailed tests of model

C / Co

Page 20: MotorMax ForceMax SpeedMax Power ROTARY ROTARY Flagellar motor (8 units, E. coli) 2400 pN nm 95 pN 300 Hz 35 m/s 2000 pN nm @ 150 Hz 1.9 x 10 6 pN nm

Resurrection

Page 21: MotorMax ForceMax SpeedMax Power ROTARY ROTARY Flagellar motor (8 units, E. coli) 2400 pN nm 95 pN 300 Hz 35 m/s 2000 pN nm @ 150 Hz 1.9 x 10 6 pN nm

resurrection

steady-state expression

One motor can contain at least 11 stators

Page 22: MotorMax ForceMax SpeedMax Power ROTARY ROTARY Flagellar motor (8 units, E. coli) 2400 pN nm 95 pN 300 Hz 35 m/s 2000 pN nm @ 150 Hz 1.9 x 10 6 pN nm

1 m bead

0.3 m bead

Page 23: MotorMax ForceMax SpeedMax Power ROTARY ROTARY Flagellar motor (8 units, E. coli) 2400 pN nm 95 pN 300 Hz 35 m/s 2000 pN nm @ 150 Hz 1.9 x 10 6 pN nm

Torque-versus speed

Page 24: MotorMax ForceMax SpeedMax Power ROTARY ROTARY Flagellar motor (8 units, E. coli) 2400 pN nm 95 pN 300 Hz 35 m/s 2000 pN nm @ 150 Hz 1.9 x 10 6 pN nm

Stepping rotation

Page 25: MotorMax ForceMax SpeedMax Power ROTARY ROTARY Flagellar motor (8 units, E. coli) 2400 pN nm 95 pN 300 Hz 35 m/s 2000 pN nm @ 150 Hz 1.9 x 10 6 pN nm

Speed control for step detection using sodium-driven chimaera

pmf or smf = Vm + kT/e ln (Cin/Cout)

Page 26: MotorMax ForceMax SpeedMax Power ROTARY ROTARY Flagellar motor (8 units, E. coli) 2400 pN nm 95 pN 300 Hz 35 m/s 2000 pN nm @ 150 Hz 1.9 x 10 6 pN nm

Low numbers of stators:

Low-level induction of stator proteins

De-energization also affects stator number

Page 27: MotorMax ForceMax SpeedMax Power ROTARY ROTARY Flagellar motor (8 units, E. coli) 2400 pN nm 95 pN 300 Hz 35 m/s 2000 pN nm @ 150 Hz 1.9 x 10 6 pN nm

Slow rotation with (probably) one stator unit

Back-focal plane detection Fluorescence detection

Real speed 30x slower

Page 28: MotorMax ForceMax SpeedMax Power ROTARY ROTARY Flagellar motor (8 units, E. coli) 2400 pN nm 95 pN 300 Hz 35 m/s 2000 pN nm @ 150 Hz 1.9 x 10 6 pN nm

26 steps per revolution

Page 29: MotorMax ForceMax SpeedMax Power ROTARY ROTARY Flagellar motor (8 units, E. coli) 2400 pN nm 95 pN 300 Hz 35 m/s 2000 pN nm @ 150 Hz 1.9 x 10 6 pN nm

Kinesin, myosin II, Myosin V, F1-ATPase:

Step size is set by the track

by energy conservation based on full energization and high loads, one proton gives a max step size of ~10 degrees. Maybe there are 2 ions per step?

34-fold model refines C-ring : 25-fold model refines M ring (& C-inner)Thomas et al 2006

One ATP per step

Page 30: MotorMax ForceMax SpeedMax Power ROTARY ROTARY Flagellar motor (8 units, E. coli) 2400 pN nm 95 pN 300 Hz 35 m/s 2000 pN nm @ 150 Hz 1.9 x 10 6 pN nm

 

 

ATP

ADP+ Pi

10nm

F1

FO

H+ or Na+

Single-molecule experiments on ATP-synthase

Page 31: MotorMax ForceMax SpeedMax Power ROTARY ROTARY Flagellar motor (8 units, E. coli) 2400 pN nm 95 pN 300 Hz 35 m/s 2000 pN nm @ 150 Hz 1.9 x 10 6 pN nm
Page 32: MotorMax ForceMax SpeedMax Power ROTARY ROTARY Flagellar motor (8 units, E. coli) 2400 pN nm 95 pN 300 Hz 35 m/s 2000 pN nm @ 150 Hz 1.9 x 10 6 pN nm

F1

Biotin-avidin link to rotating handle(actin filament, beads)

His-tag link to surface

Page 33: MotorMax ForceMax SpeedMax Power ROTARY ROTARY Flagellar motor (8 units, E. coli) 2400 pN nm 95 pN 300 Hz 35 m/s 2000 pN nm @ 150 Hz 1.9 x 10 6 pN nm

0.5 micron beadsRotated by F1

Fluorescent actin filamentRotated by F1

(movie: Wolfgang Junge)

Page 34: MotorMax ForceMax SpeedMax Power ROTARY ROTARY Flagellar motor (8 units, E. coli) 2400 pN nm 95 pN 300 Hz 35 m/s 2000 pN nm @ 150 Hz 1.9 x 10 6 pN nm
Page 35: MotorMax ForceMax SpeedMax Power ROTARY ROTARY Flagellar motor (8 units, E. coli) 2400 pN nm 95 pN 300 Hz 35 m/s 2000 pN nm @ 150 Hz 1.9 x 10 6 pN nm

High [ATP] : no wait before 90° step

Medium [ATP] : ~ms before 90° step

Low [ATP] : long wait before 90° step

(ms)

All [ATP] : ~ms before 30° step

Page 36: MotorMax ForceMax SpeedMax Power ROTARY ROTARY Flagellar motor (8 units, E. coli) 2400 pN nm 95 pN 300 Hz 35 m/s 2000 pN nm @ 150 Hz 1.9 x 10 6 pN nm

Low [ATP] : - 90° step rate-limiting- exponential distribution- single step

High [ATP] : - 30° step rate-limiting- peaked distribution- double (or more) step

Page 37: MotorMax ForceMax SpeedMax Power ROTARY ROTARY Flagellar motor (8 units, E. coli) 2400 pN nm 95 pN 300 Hz 35 m/s 2000 pN nm @ 150 Hz 1.9 x 10 6 pN nm
Page 38: MotorMax ForceMax SpeedMax Power ROTARY ROTARY Flagellar motor (8 units, E. coli) 2400 pN nm 95 pN 300 Hz 35 m/s 2000 pN nm @ 150 Hz 1.9 x 10 6 pN nm
Page 39: MotorMax ForceMax SpeedMax Power ROTARY ROTARY Flagellar motor (8 units, E. coli) 2400 pN nm 95 pN 300 Hz 35 m/s 2000 pN nm @ 150 Hz 1.9 x 10 6 pN nm

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