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For a circular domain of radius $R_0$ with line tension $\gamma$, the stiffness of a real polar sine or cosine amplitude with integer angular harmonic $m$ is $K_m=\pi \gamma (m^2-1)/R_0$, when the enclosed area is fixed. The equipartition theorem gives $⟨u_m^2⟩=k_{B}T{R}_0/[\pi \gamma (m^2-1)]$ for $m\ge 2$. A positive-frequency complex Fourier mode coefficient has half this variance. The uniform mode is forbidden and $m=1$ is translation.

For a tension-dominated cylindrical lipid vesicle of fixed length $L$ and volume, a real axisymmetric sine amplitude $u_q$ has stiffness $K_q=\pi \sigma L[(q{R}_0{)}^2-1]/R_0$. Fixed volume requires a mean-radius shift $-u_q^2/(4R_0)$. Stable modes have $⟨u_q^2⟩=k_{B}T/K_q$ by the equipartition theorem. Modes with $q{R}_0<1$ have the Rayleigh–Plateau instability, rather than a negative equilibrium variance. Bending modifies very short wavelengths and near-marginal modes.

Choose real eigenfunctions with ${\int }_0^L{W}_n{W}_{m}dx={\delta }_{nm}$. For positive modes, self-adjointness and integration by parts diagonalize the energy:At temperature $T$, the canonical ensemble is a product of centered Gaussian distributions. The equipartition theorem, with Boltzmann constant $k_B$, givesThe resulting thermal covariance of an elastic filament isFor an unnormalized eigenfunction of squared L2 norm $N_n$, divide its summand by $N_n$. This normalization factor cannot be absorbed silently into the modal variance.

For clamped boundary conditions at both ends, the inverse of $d^4/d{x}^4$ has Green function, for $x\le y$,It is cubic on each side of $y$, satisfies the four clamped conditions, has continuous first two derivatives, and has unit jump in its third derivative. Thus ${\partial }_x^{4}G=\delta (x-y)$, and its eigenfunction expansion is the sum above. In particular,These finite variances require removal of every zero-energy filament mode. An unconstrained free-free elastic filament can translate and tilt at no energy cost; a torqued-torqued elastic filament can translate. Their unrestricted Boltzmann distributions are not normalizable, so the full displacement variance is undefined. Fix those rigid degrees of freedom before applying the positive-mode formula.

For spatially varying tension in filament bending, keep the derivative of the filament tension as well as the curvature term. The first variation isTherefore the Euler-Lagrange equation and fluctuation operator areFor real, sufficiently smooth $\sigma$, the boundary form is the bending boundary form minus $[\sigma (\overline{u}{v}^{\prime }-\overline{u^{\prime }}v){]}_0^L$. Because $\sigma$ vanishes at both ends, the four self-adjoint endpoint conditions for filament bending still apply. The natural endpoint force also reduces there to the bending shear term. Thus $K_{\sigma }$ is a self-adjoint fourth-order scalar differential operator on the same chosen domain, with compact resolvent.

Choose a real orthonormal basis ${\psi }_n$ of eigenfunctions, $K_{\sigma }{\psi }_n={\lambda }_n{\psi }_n$, and write $h={\sum }_n{b}_n{\psi }_n$. Using the endpoint conditions in integration by parts givesThe equipartition theorem now gives, on the strictly positive subspace,This is a formal modal construction; no explicit eigenfunctions are needed. Nonnegative filament tension makes the energy nonnegative. Any surviving zero-energy filament modes must again be fixed. If signed $\sigma$ permits compression, self-adjointness still holds but does not guarantee a canonical ensemble: sufficiently strong compression can create negative eigenvalues and Euler buckling of an elastic filament. For instance, on $A=L=1$, take $\sigma =-Cx(1-x)$ and the clamped trial function $h=x^2(1-x{)}^2$. Thenso the energy is negative when $C>252$, despite $\sigma (0)=\sigma (1)=0$. The equipartition theorem requires a stable positive quadratic energy, not merely a real modal spectrum.

Take a long periodic section of length $L$, or neglect end effects and choose an integer number of wavelengths. The amplitude $u_q$ here is a single real sine amplitude, fixing the normalization of the thermal membrane undulation spectrum. For a lipid vesicle with an axisymmetric membrane deformation, conservation of the enclosed volume givesThe mean radius therefore decreases at second order. This adjustment is essential: fixing the mean radius instead of the volume would miss the unstable term in the fixed-volume capillary spectrum of a cylindrical membrane.

The surface area of an axisymmetric graph is $\mathcal{S}=2\pi \int R\sqrt{1+R_z^2}dz$. Expanding for $|u_q|\ll R_0$ and $q|u_q|\ll 1$ givesMultiplication by the membrane tension yields a quadratic potential energy $\Delta E=K_q{u}_q^2/2$, whereFor $q{R}_0>1$, the equipartition theorem gives the stable-mode fixed-volume capillary spectrum of a cylindrical membraneHere $k_B$ is the Boltzmann constant and $T$ is the temperature. The cosine amplitude has the same variance and is an independent real coordinate at quadratic order. If instead $R-{\rho }_0={\sum }_{q\ne 0}{a}_q{e}^{iqz}$ with $a_{-q}=a_q^{\ast }$, the corresponding positive-$q$ complex coefficient has $⟨|a_q{|}^2⟩=k_{B}T{R}_0/[2\pi \sigma L((q{R}_0{)}^2-1)]$. This is the same thermal membrane undulation spectrum in a different Fourier mode normalization.

For $q{R}_0<1$, the cylinder is unstable rather than having a negative fluctuation variance. The quadratic surface area change is negative: a sufficiently long-wavelength modulation reduces area while retaining volume. This is the Rayleigh–Plateau instability, expressed here as pearling of a tension-dominated lipid vesicle. A cylinder supporting such modes has no unconstrained harmonic thermal equilibrium about the uniform state. The $q=0$ uniform radius change is forbidden by fixed volume, and $q{R}_0=1$ is marginal in the tension-only approximation. The finite length and endpoint constraints determine which nonzero Fourier modes are allowed.

The large membrane tension assumption controls modes with $q{R}_0$ of order one. For large $q$ the omitted membrane bending modulus contribution grows approximately as $\kappa q^4$ and overtakes the tension term when $\kappa q^2\gtrsim \sigma$. Thus the displayed tension-only spectrum requires $q\ll \sqrt{\sigma /\kappa }$ as well as small amplitudes and a positive stiffness. Close enough to the marginal wavelength, bending corrections must also be retained; their relative scale at $q{R}_0$ of order one is $\kappa /(\sigma R_0^2)$.

For the circular phase boundary in a lipid bilayer, the energy is its perimeter times the line tension $\gamma$. Let $\phi$ be the polar angle and write $r(\phi )={\rho }_0+u_m\sin (m\phi )$, where the periodicity requires an integer $m=q{R}_0\ge 1$. Fixed enclosed area impliesThe perimeter, including the mean-radius change, isThus the fixed-area capillary spectrum of a circular boundary, again for a single real sine or cosine amplitude, isThe complex Fourier mode coefficient convention again divides this result by two. The mode $m=0$ is excluded by the fixed-area constraint, so the negative-stiffness interval $0<m<1$ is not an allowed mode of a closed circular boundary.

At $q{R}_0=1$, the zero stiffness is the translation mode of a circular boundary, not a shape instability. Displacing the centre by a small distance $h$ changes the radius to first order by $h\cos \phi$ or $h\sin \phi$ while leaving area and perimeter unchanged. For example, an exactly translated circle hasThe higher harmonics complete a true translation mode of a circular boundary. The $m=1$ denominator therefore represents free centre motion: an unconfined domain's centre has no restoring force or finite equilibrium variance in an infinite membrane. Radius fluctuations about a recentered domain omit this mode; external confinement would give it a separate restoring stiffness.

For one small transverse displacement, the elastic energy of the elastic filament iswhere $A$ is the filament bending modulus. Its first variation isThe clamped boundary conditions remove the left boundary terms. A force-free and torque-free tip permits independent $\delta h(L)$ and $\delta h^{\prime }(L)$, so the natural boundary conditions for a free endpoint are $h^{\prime \prime }(L)=h^{\prime \prime \prime }(L)=0$: zero bending moment and transverse shear.

There is an important distinction between an energy minimum and a normal mode. The unloaded higher-order Euler-Lagrange equation is $h^{\prime \prime \prime \prime }=0$, whose only solution with these four boundary conditions is $h=0$. A general fluctuating shape is a sum of modes, not one of the stated sinusoidal/hyperbolic functions. To obtain the clamped--free bending modes, extremize the bending Rayleigh quotient, or equivalently $U[W]-\frac{1}{2}A{k}^4\int W^{2}dx$ with a fixed norm. Its Euler-Lagrange equation isThis is the constrained variational characterization of bending modes. Its four characteristic roots are $\pm k,\pm ik$. Clamping gives $F=-D$ and $E=-B$, henceThe free-tip conditions reduce toIts determinant is $-2(1+\cos q\cosh q)$, so nontrivial modes requireNo zero mode exists, since a cubic satisfying the homogeneous clamp/free conditions is zero. The clamped-free bending spectrum starts with $q_1\in (\pi /2,2)$; bisection or Newton iteration gives $q_1\simeq 1.875104$, so $k_1\simeq 1.875104/L$. Choosing $D=N(\sin q_n+\sinh q_n)$ and $B=-N(\cos q_n+\cosh q_n)$ givesThe second boundary condition follows from the root equation, and $N$ is arbitrary until a normalization is chosen.

The bending operator with these boundary conditions is positive and self-adjoint. Twice integrating by parts givesSymmetry implies orthogonality when $n\ne m$. Expand $h(x)={\sum }_n{a}_n{W}_n(x)$ and use $I_n=L^{-1}{\int }_0^L{W}_n^{2}dx$. Thenby the equipartition theorem. The supplied endpoint identity therefore yields the thermal bending fluctuations of a clamped filament:To evaluate the sum, apply a tip force $f$ and minimize $U-fh(L)$. The modified free-end conditions are $h^{\prime \prime }(L)=0$ and $-A{h}^{\prime \prime \prime }(L)=f$, and $h^{\prime \prime \prime \prime }=0$ in the interior. Integration gives $h(x)=f{x}^2(3L-x)/(6A)$, so the tip-force compliance of a cantilever is $L^3/(3A)$. On the other hand, minimizing the modal energy minus $f{\sum }_n{a}_n{W}_n(L)$ gives $a_n=f{W}_n(L)/(AL{k}_n^4{I}_n)$ and henceComparison evaluates the fourth inverse-power sum of the cantilever spectrum without truncating the modes:The boxed variance is for the specified single transverse direction. An independent equilibrium check follows by differentiating the Gaussian partition function with respect to $f$: $\partial ⟨h(L)⟩/\partial f=⟨h(L{)}^2⟩/(k_{B}T)$ at zero load, reproducing the same result from the static compliance. Keeping only the first normal mode gives about $0.32356k_{B}T{L}^3/A$, roughly $97.1\%$ of the exact variance. With two independent equivalent transverse directions, their summed variance is twice the boxed result. The small-slope model requires $k_{B}TL/A\ll 1$, or $L$ small compared with the usual three-dimensional persistence length $A/(k_{B}T)$.

For $q\ne 0$ and $\alpha >0$, each Fourier mode is an Ornstein-Uhlenbeck process with decay rate $r_q=\alpha q^2$. Its explicit Ornstein-Uhlenbeck solution isso the Itô isometry givesTo identify the Overdamped Langevin dynamics, choose a friction ${\zeta }_q>0$, a harmonic oscillator potential $V_q={\zeta }_q\alpha q^2{h}_q^2/2$, and force noise $f_q={\zeta }_q{\eta }_q$. The fluctuation-dissipation relation for a Langevin particle then defines $k_B{T}_{\mathrm{eff}}={\zeta }_q{\sigma }^2/2$. Its Boltzmann distribution isThis is the stationary spectrum of a linear fluctuating interface, also following from the equipartition theorem. The numerical prefactor uses exactly the mode-noise normalization given in the paper. The nonzero modes can therefore have a stationary Gaussian distribution even though the unpinned zero mode cannot.

For a nearly planar membrane, the quadratic Helfrich energy of a Fourier mode isThe equipartition theorem and $A-A_{\mathrm{proj}}=\int |\nabla h{|}^2{d}^{2}x/2$ giveUsing the stated molecular and system-size cutoffs, $q_{\max }=\pi /a$ and $q_{\min }=\pi /\sqrt{A}$, gives

In the small-slope approximation the worm-like chain bending energy isThe clamped end obeys $h(0)=h_x(0)=0$, while a force-free and torque-free tip has the natural conditions $h_{xx}(L)=h_{xxx}(L)=0$. A static transverse tip force $f$ produces the cantilever compliance $h(L)/f=L^3/(3A)$. The equipartition theorem, or equivalently the static fluctuation--response relation, therefore gives the exact tip variance

For the dynamics, resistive-force theory gives the transverse drag per lengthat logarithmic accuracy and the stochastic beam equationLet ${\varphi }_n$ be clamped--free bending modes with $q_{n}L={\alpha }_n$, whereWriting $N_n={\int }_0^L{\varphi }_n^{2}dx$, independent thermally driven modes giveThe first mode contains almost all of the tip variance, so

For a microtubule, $A=k_{B}T{L}_p$. With $L=10\mu \mathrm{m}$ and $L_p=3\mathrm{mm}$,Taking water at room temperature and $a=0.5\mu \mathrm{m}$ gives ${\zeta }_{\perp }\simeq 3.6\times 10^{-3}\mathrm{Pa}\mathrm{s}$ andwith an order-one uncertainty from the logarithmic slender-body drag approximation.

Ifthen $e^{\beta a}$ is large and the wall correction is exponentially small:This is exactly the equipartition theorem result from the two kinetic and two harmonic-potential quadratic terms.

If $a\ll k_{B}T$, useThenThe leading term is the kinetic energy. The particle is nearly uniform in the disk, so $⟨r^2⟩\to R^2/2$ and $⟨V⟩\to \kappa R^2/4$, much smaller than $k_{B}T$. The hard wall therefore suppresses the positional equipartition contribution, as in the hard-wall correction to harmonic equipartition.

PutThe thermodynamic derivatives of the canonical partition function giveThe two momentum coordinates are unrestricted quadratic degrees of freedom, so the equipartition theorem gives mean kinetic energy $1/\beta$. HenceThe same result follows from $⟨V⟩=-(\kappa /\beta ){\partial }_{\kappa }\log Z_1$.