Chapter 8 – Theoretical Biophysics  389

a

By treating each link as a position vector, and neglecting possible consequences

of interference between different parts of the chain, derive an expression for <x2>,

the mean square end-​to-​end distance.

b

Evaluate this expression in the limits that the contour length is much greater, and

much less, than b, and comment on both results.

8.6

A certain linear protein molecule was modeled as an ideal chain with 500 chain

segments each equal to the average alpha-​carbon spacing for a single amino acid

carrying a charge of ±q at either end of each amino acid subunit where q is the unitary

electron charge. What is the end-​to-​end length relative to its rms unperturbed length

parallel to an E-​field, of 12,000 V cm−1?

8.7

A protein exists in just two stable conformations, 1 and 2, and undergoes reversible

transitions between them with rate constants k12 and k21, respectively, such that the

free energy difference between the two states is ΔG.

a

Use detailed balance to estimate the probability that the system is in state 1 at

a given time t, assuming that initially the molecule is in state 1. A diferent pro­

tein undergoes two irreversible transitions in conformation, 1 –​ 2 and 2 –​ 3, rate

constants k12 and k23, respectively.

b

What is the likelihood that a given molecule starting in state 1 initially will be in

state 3 at time t?

8.8

In an experiment, the total bending free energy of a liposome was measured at ~200

kBT and the polar heads tightly packed into an area equivalent to a circle of radius of

0.2 nm each, with the length from the tip of the head group to the end of the hydro­

phobic tail measured at 2.5 nm. Estimate the free energy in Joules per phospholipid

molecule required to double the area occupied by a single head group.

8.9

A virus was fluorescently labeled and monitored in a living cell at consecutive sam­

pling times of interval 33 ms up to 1 s. The rms displacement was calculated for two

such viral particles, giving values of [61, 75, 81, 95, 107, 112, 128, 131, 158, 167, 181,

176, 177, 182, 183, 178, 177, 180, 182, 184, 181, 179, 180, 178, 180, 182] nm and [59,

65, 66, 60, 64, 63, 58, 62, 63, 61, 64, 60, 59, 64, 62, 65, 61, 60, 63, 66, 62, 58, 60, 57,

61, 62] nm. What might this indicate about the virus diffusion?

8.10 A series of first-​order biochemical reactions of molecules 1, 2, 3,…, n reacted irrevers­

ibly as 1 2 3 n with rate constants k1, k2, k3,…,kn, respectively.

a

What is the overall rate constant of the process 1 n?

b

What happens to this overall rate if the time scales for formation of one of the

intermediate molecules are significantly lower than the others?

c

What implications does this have for the local concentrations of the other inter­

mediate molecules formed? (Hint: such a slow-​forming intermediate transition is

often referred to as the “rate-​limiting step.”)

8.11 Write down the Brownian diffusion equation for numbers of molecules n per unit

length in x that spread out by diffusion after a time t.

a

Show that a solution exists of the form n(x,t) =​ αt½exp(−x2/​(4Dt)) and determine

the normalization constant a if there are ntot molecules in total present.

b

Calculate the mean expected values for <x> and <x2>, and sketch the solution of

the latter for several values of t > 0. What does this imply for the location of the

particles initially?

The longest cells in a human body are neurons running from the spinal cord to

the feet, roughly shaped as a tube of circular cross-​section with a tube length

of ~1 m and a diameter of ~1 μm with a small localized cell body at one end

that contains the nucleus. Neurotransmitter molecules are synthesized in the

cell body but are required at the far end of the neuron. When neurotransmitter

molecules reach the far end of the neuron, they are subsequently removed by a

continuous biological process at a rate that maintains the mean concentration

in the cell body at 1 mM.

c

If the diffusion coefficient is ~1000 μm2s−1, estimate how many neurotransmitter

molecules reach the end of the cell a second due to diffusion.