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Propulsion and Evolution of Algae. R E Goldstein DAMTP Cambridge. ?. The Size-Complexity Relation. Amoebas, Ciliates, Brown Seaweeds Green Algae and Plants Red Seaweeds Fungi Animals. Bell & Mooers (1997) Bonner (2004). Volvox. Phil. Trans . Roy. Soc. 22 , 509-518 (1700). (1758). - PowerPoint PPT Presentation
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Propulsion and Evolution of Algae
R E GoldsteinDAMTPCambridge
The Size-Complexity Relation
Amoebas, Ciliates, Brown SeaweedsGreen Algae and PlantsRed SeaweedsFungiAnimals
Bell & Mooers (1997)Bonner (2004)?
Volvox
Phil. Trans. Roy. Soc. 22, 509-518 (1700)
(1758)
A Family PortraitChlamydomonas reinhardtii
Volvox carteri
Gonium pectorale Eudorina elegans
Pleodorina californica
Volvox aureus
daughter colonies somatic cellsGerm-soma differentiation
Altruism, apoptosis
The Diffusional Bottleneck
Organism radius R
Cu
rren
ts
RCDI
r
RCC
4
1
d
Diffusion to an absorbing sphere
PO42- and O2 estimates yield
bottleneck radius ~50-200 m(~Pleodorina, start of germ-somadifferentiation)
DCRb
Metabolic requirements scale with surface somatic cells
2m 4 RI
Advection & Diffusion
If U=10 m/s, L=10 m, Pe ~ 10-1
At the scale of an individual cell, diffusion dominates advection.
The opposite holds for multicellularity…
D
UL
t
tPe
advection
diffusion
If a fluid has a typical velocity U, varying on a length scale L, with a molecular species of diffusion constant D. Then there are two times:
D
Lt
U
Lt
diffusion
advection
2
We define the Péclet number as the ratio:
Microscopy & Micromanipulation
motorized microscope stage
micro-manipulator
micro-manipulator
Stirring by Volvox carteri P
seu
do-
da
rkfie
ld (
4x o
bje
ctiv
e, P
h4
rin
g)
To
ols
of t
he
tra
de –
mic
ropi
pet
te
pre
pa
ratio
n
1 mm
A Closer ViewF
luo
resc
enc
e
Fluid Velocities During Life Cycle
Hatch Division Daughter Pre-Hatch
This is “Life at High Péclet Numbers”
D
RuPe max2
Metabolite Exchange
Flagella Beating/Symmetry
(2000 frames/s background subtraction)
Noisy Synchronization
Experimental methods:• Micropipette manipulation with a rotating stage for precise alignment• Up to 2000 frames/sec• Long time series (50,000 beats or more)• Can impose external fluid flow
Micropipette
Cell body
Frame-subtraction
22
11
dt
ddt
d
)](cos[A)(
)](cos[A)(
222
111
ttS
ttS
“Phase oscillator” model used in e.g. circadian rhythms, etc.
t2121
strokes offlagella
amplitudes “phases”or angles
naturalfrequencies
Historical Background• R. Kamiya and E. Hasegawa [Exp. Cell. Res. (‘87)] (cell models – demembranated) intrinsically different frequencies of two flagella
• U. Rüffer and W. Nultsch [Cell Motil. (‘87,’90,’91,’98)] short observations (50-100 beats at a time, 1-2 sec.) truly heroic – hand drawing from videos synchronization, small phase shift, occasional “slips”
Without coupling, the phase difference simply grows in time
So, is this seen?
Key issue:control ofphototaxis
A Phase Slip
Dynamics of Phase Slips (Both Directions!)
Drifts and Slips are Controlled by the Cell
frequency (arb)
Po
wer
sp
ectr
um
“Random” Swimming of Chlamydomonas reinhardtii
Red light illumination – no phototactic cues45 s. track – note many changes of directionVolume explored is ~1 mm3 very far from chamber walls
Geometry of Turning
beat
rad4.0
beats16~
2
s
beats10-5
rad/s 4-2
s1-0.5
beats5025
driftT 90rad21driftT
Turning angle (degrees)
~100o
90
Pro
bab
ility
(a
ngle
)
Chlamy w/single flagellum,rotating near a surface
Angle per beat -
Frequency difference -
“Drift” duration-
Angular velocity
Angular change
Walzing Volvox: Orbiting “Bound State”
Dual Views
Dominant physics: downwardgravitational force on the colony,producing recirculating flows.
Fluid flow produced by a point force near a wall: solved exactly by J.R. Blake (1971)
The Minuet Bound State
Numerical solution of a model:Based on hovering, negativelybuoyant, bottom-heavy swimmers.
Bottom-heaviness confers stability.
Side view
Chamber bottom
Marco PolinIdan TuvalKyriacos LeptosKnut Drescher
Sujoy Ganguly
Cristian Solari
Timothy J. PedleyTakuji Ishikawa
Jerry P. Gollub
www.damtp.cam.ac.uk/user/gold
Our Team