104 3.6  Basic Fluorescence Microscopy Illumination Modes

characterized by fast diffusion of one more molecular component but often slow reaction

events with other components (see Chapter 8). Thus, the time scale for the entire process may

be substantially longer than the millisecond diffusional time scale. To overcome this issue,

individual millisecond image frames may be obtained discontinuously, that is, using strobing.

In this way, the limited fluorescence emission photon budget of a single fluorescent protein

molecule may be used over substantially longer time scales than just a few tens of milliseconds

to gain insight into the dynamics of several different processes inside the cell while still enab­

ling unblurred fluorophore detection on individual images frames. A technical issue in regard

to strobing is the bandwidth of the shuttering mechanism used to turn the laser excitation

on and off. Typical mechanical shutters are limited to a bandwidth of ~100 Hz, and so the

minimum sampling time window that can be accommodated is ~10 ms, which is too high for

millisecond imaging. Alternative faster shuttering can be implemented directly through elec­

tronic modulation of the laser power output on some devices or utilizing acousto-​optic-​based

technology for shuttering, for example, an acousto-​optic modulator that can be shuttered at

>MHz bandwidth using similar physics principles to AODs (see Chapter 6).

KEY POINT 3.5

In fluorescence imaging at the molecular scale, many single-​particle tracks are

truncated, often because of rapid fluorophore photobleaching. Strobing can permit the

limited photon emission budget to be stretched out over larger time scales to probe

slower biological processes.

Worked Case Example 3.3: Slimfield

A Slimfield microscope comprised a 488 nm wavelength laser of beam width 0.1 mm

directed into the back aperture of a high numerical aperture objective lens of NA 1.45, focal

length 2 mm, to image a single GFP molecule at room temperature, peak fluorescence

wavelength ~505 nm, tagged to a cell membrane protein such that the protein is integrated

into the phospholipid membrane. The membrane viscosity was measured separately to be

~100 cP, but the GFP molecule itself sits just outside the membrane in the cytoplasm, whose

viscosity was closer to 1 cP. The cell membrane is in contact with the microscope coverslip

and can be considered reasonably planar over a circle of diameter ~5 µm.

a If the frictional drag coefficient of the membrane integrated protein is 1.1 × 10-​7 kg/​s,

what is the maximum sampling time per image frame which will allow you to observe

the GFP unblurred? Before the protein integrates into the membrane, it must first diffuse

through the 3D volume of the cytoplasm. If you wish to monitor this cytoplasmic diffusion

as molecules come into focus before they integrate into the membrane, what is the max­

imum sampling time that could be used?

b In a separate experiment, the average time that a single GFP molecule fluoresces

before photobleaching using identical Slimfield imaging conditions as that for measuring

the cytoplasmic diffusion was measured to be 30 ms. If the membrane protein is integrated

into a lipid raft region of the cell membrane whose effective diameter is 200 nm, do you

think you will be able to also monitor the diffusion of the membrane integrated protein as

it moves from the center to the edge of a raft?

Answers

a

The PSF width for imaging a GFP molecule is given by Equation 3.54 of w=​0.61λ/​

NA. Here λ relates to the fluorescence emission, not the excitation, so using the

characteristic peak value given indicates that:

w =​ 0.61 × (505 × 10-​9)/​1.45 = ​212 × 10–​9 m (i.e. 212 nm)

KEY BIOLOGICAL

APPLICATIONS:

DIFFERENT

FLUORESCENCE

MICROSCOPY MODES

Investigating cell membrane

processes in live cells; Probing

rapid processes in the cell cyto­

plasm of live cells; Rendering

axial as well as lateral localization

information for processes and

components inside cells.