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# A CHANDRA SEARCH FOR X-RAY JETS IN REDSHIFT 6 QUASARS

## 1 INTRODUCTION

The sub-arcsecond angular resolution of the *Chandra* X-ray Observatory has proven to have unique capability to resolve jets and hot-spots in quasars and radio sources. This was shown in the very first pointed observation (Schwartz et al., 2000; Chartas et al., 2000), at the z=0.652 quasar PKS 0637–752 for which Schwartz et al. (2000) showed that the X-rays could not plausibly arise from the synchrotron mechanism. To explain the X-ray emission, Tavecchio et al. (2000), and Celotti et al. (2001) suggested that bulk relativistic motion of the jet would allow the X-rays to be produced by inverse Compton (IC) radiation from the cosmic microwave background (CMB), while allowing the magnetic fields and relativistic electrons to be near equipartition in the jet rest frame. Schwartz (2001, 2002) noted that if observed X-rays were produced by IC/CMB, whether or not relativistically beamed, then such a source would be seen at any larger redshift with the same surface brightness, and would be resolved and detected by *Chandra*.

Fan et al. (2001) have discovered the three most distant quasars known, at redshifts 6.28, 5.99, and 5.82. A Director’s Discretionary Program (cf. http://cxc.harvard.edu/DDT/DDTobs\_info.html) proposed by Brandt was carried out on 29 January, 2002 to obtain “snapshot” data for immediate public use in planning further studies of distant quasars. We have used these data to search for X-ray jets associated with those quasars.

## 2 OBSERVATIONS OF THREE SDSS QUASARS AT REDSHIFT 6

The observations were carried out in the standard 6-chip ACIS-S imaging mode, with a 3.2s frame time, and events telemetered in the 3$`\times `$3 faint mode format. We have selected for analysis only the data from S3, with ASCA grades 0,2,3,4,6, and energy between 0.5 and 7 keV. At lower energy the quantum efficiency and energy response is increasingly uncertain, and at higher energies the background is increasing steeply.

Table 1 summarizes the observations. The observing time is taken from the number of frames, times the 3.2s frame time. For the core, we tabulate the measured counts in a 1$`\stackrel{}{\mathrm{.}}`$23 radius about the quasar position (5 pixel diameter), giving $``$ 95% encircled energy fraction at 1.5 keV (Jerius, et al., 2000). We convert from a measured counting rate in the 0.5–7 keV band to an unabsorbed, measured flux in that band assuming a power law spectrum of energy index 0.7, and the galactic hydrogen column density. This is approximately 6.0$`\times `$10-12 ergs cm-2 s-1 per count s-1. The factor changes about $`\pm `$ 1.5% for the range of column densities of the three objects, 2.1 to 4.4 $`\times `$1020cm-2, and changes by about $`\pm `$ 20% for indices from 0.4 to 1.2. Poisson statistics on the few detected counts dominate the uncertainty. The final two columns convert to rest frame luminosity in the 2–10 keV band111We use $`H_0=65\mathrm{km}\mathrm{s}^1\mathrm{Mpc}^1`$ and a flat universe with $`\mathrm{\Omega }_0=0.3`$, $`\mathrm{\Omega }_\mathrm{\Lambda }=0.7`$, and q0 = -0.55 throughout., including the K-correction for the assumed spectrum.

Figure 1 shows the detection of each quasar. The plus sign gives the true position of the quasar (Fan et al., 2001). The X-ray centroids are offset by 0$`\stackrel{}{\mathrm{.}}`$65, 0$`\stackrel{}{\mathrm{.}}`$76, and 0$`\stackrel{}{\mathrm{.}}`$66, respectively for SDSS 0836, SDSS 1030, and SDSS 1306. These are within the astrometric performance of *Chandra* (cf. http://cxc.harvard.edu/cal/ASPECT/celmon/). Each quasar is clearly detected. The background rates are determined from large rectangular regions to be 2.04, 2.56, and 2.92 $`\times `$10-3 counts per pixel, respectively, so there is a negligible background correction of 0.04 to 0.05 counts, which we do not make. In the 19 pixel source extraction circles, the probability is less than 2.6$`\times `$10-5 of getting 3 or more counts, so all the detections are highly significant. (For a serendipitous source anywhere in the field, one only needs 4 counts to have less than a 1% probability of a chance occurrence.) SDSS 0836 and SDSS 1306 are clearly consistent with being point sources.

The 6 photons from SDSS 1030+0524 are highly unlikely to be from a single point source. They are all further than 0$`\stackrel{}{\mathrm{.}}`$68 from their centroid. That distance is the 79% encircled energy radius, so the probability is 8.6$`\times \mathrm{\hspace{0.17em}10}^5`$ that all six photons would fall outside this radius. There is a 4.7% chance that one is due to background, but less than 0.13% chance that two or more are background counts. If we discard the most distant photon and recompute the X-ray centroid, one falls at the 55% encircled radius and the other four are still outside the 80% radius, and the probability of at least four so distant from the centroid is only 6.7$`\times 10^3`$. Therefore either the source is extended, over about a 2″ diameter, or it is really two or more point sources. As discussed by Wyithe & Loeb (2002a, b), there is a probability between 7% and 30% that any quasar at z=6 is gravitationally lensed, and this could be confirmed with a followup observation of $`>`$50 ks.

Figure 2 shows the region surrounding each quasar. For the cosmological parameters used here, 1$`\mathrm{}`$ corresponds to 6.26, 6.00 and 6.16 kpc for SDSS 0836, SDSS 1030, and SDSS 1306, respectively. The larger circles indicate a 100 kpc projected distance from each quasar. There are no significant concentrations of photons anywhere within these circles. In the annulus we expect 6.8, 9.3, and 10.0 background counts and observe 6, 14, and 8, respectively.

SDSS 1306+0356 has a significant X-ray feature 23$`\stackrel{}{\mathrm{.}}`$3 to the NE of the quasar, at a distance 143 kpc projected in the plane of the sky. There are 7 counts in a box 5$`\mathrm{}\times `$2$`\mathrm{}`$ which points toward the quasar. The significance of these as an X-ray source is beyond doubt. We can arbitrarily place this box to include at least one count. Since there are only 0.13 background counts expected, the chance of getting 6 additional counts in a 10 arcsec2 box is only 5.8$`\times `$10-9, or only 1.7$`\times `$10-4 of occurring in any 10 arcsec2 box within 5$`\mathrm{}`$ of the quasar. There is no visible object at this position in the digital sky survey. The FIRST radio survey gives an upper limit of 0.93 mJy at 1.4 GHz at this position (White et al., 1997). Interpretation as an IC/CMB X-ray jet is entirely reasonable, as we discuss below; however, there is no specific evidence requiring such interpretation. Arguments as given above for the core of SDSS 1030 rule out a single point source, with or without attributing any of the 7 counts to background. The spatial distribution of counts cannot rule out two point sources and a background count. Giacconi et al. (2001) give about 200 sources deg-2 above the flux of 5$`\times `$10-15 observed from the jet, so there is a 5% chance of one such source falling within 1$`\mathrm{}`$ of the quasar. The chance occurrence of an extended source at this flux level is at least 10 times smaller, $`<`$0.5% (see Bauer et al. (2002)). Because a longer X-ray observation is needed to obtain definitive information, and because we cannot rule out the possibility of foreground sources correlated with each other, we do not attempt a more exhaustive discussion of the probabilities. As a newly discovered X-ray source, we designate this as CXOU 130609.31+035643.5.

In Table 1 we set a limit to the flux of putative jets in SDSS 0836 or SDSS 1030 by noting that we do not find 3 or more photons in any similar 10 arcsec2 box pointing toward the core. If there were a jet from which we expected 6.3 or more counts, then there would be less than a 5% probability to fail to detect 3 or more, and we adopt this as our limit.

Because of the small numbers of photons, we need to give careful consideration to systematic effects. Figure 3 shows that the background was steady during these observations, and the arrival times of the quasar and jet counts are consistent with being uniform throughout the observations. No two counts were registered from the same physical pixel.

To investigate any peculiarities in the distribution of photon energies, we perform a KS test of the energy distributions of the photons from each quasar and from the jet feature against the core spectrum of 3C 273. We might expect them *a priori* to have a non-thermal spectrum of energy index roughly $`\alpha `$0.7. Figure 4 shows the cumulative distributions. The solid heavy line is the core spectrum of 3C 273, determined from 3063 photons in the readout streak in OBSID 1712 (Marshall et al., 2001). If SDSS 0836, SDSS 1030, SDSS 1306, and the jet all had the same spectrum as 3C 273, we would expect deviations as large as actually observed 98%, 16%, 96%, and 10% of the time, respectively. Thus we cannot reject this hypothesis for any of them. The crosses plot the distribution of 763 background photons from a source free region of the SDSS 1306 observation. Such large deviations as observed would be expected only 0.3%, 5.6%, 2.8%, and 90% of the time, for the same respective sources. Thus the quasar core spectra are very distinct from that of the background, while the putative jet spectrum is consistent with the harder background spectral shape.

We compute $`\alpha _{\mathrm{ox}}`$ according to the original definition (Tananbaum et al., 1979),

$$\alpha _{\mathrm{ox}}=\mathrm{log}(f_x/f_o)/\mathrm{log}(\nu _x/\nu _o),$$
(1)
where $`f_x`$ is the X-ray flux density at 2 keV, $`\nu _x`$= 4.8$`\times `$1017 Hz, and $`f_o`$ is the optical flux density at 2500 Å, $`\nu _o`$=1.2$`\times `$1015 Hz. The flux densities are computed in the rest frame. We take the AB1450 magnitudes from Fan et al. (2001) and extrapolate from 1450Å to 2500Å assuming a power law flux $`\nu ^{0.5}`$ (Richstone & Schmidt, 1980; Vanden Berk et al., 2001). We find $`\alpha _{\mathrm{ox}}`$ of 1.66, 1.79, and 1.65, respectively, for SDSS 0836, SDSS 1030, and SDSS 1306. These values are consistent with the range found by Vignali et al. (2001) for *Chandra* quasars at z$`>`$4, but about 0.1 larger numerically than the mean found by Kaspi, Brandt, & Schneider (2000) for *ROSAT* z$`>`$4 quasars. We note that an extraction circle larger than 25$`\mathrm{}`$ applied to SDSS 1306+0356 would lead to a decrease in $`\alpha _{\mathrm{ox}}`$ by 0.06, due to inclusion of flux from CXOU 130609.31+035643.5.

## 3 CAN CXOU 130609.31 BE AN X-RAY JET FROM SDSS 130608.26?

The feature discovered by *Chandra* to the NE of SDSS 1306 can be explained as a jet emitting by inverse Compton scattering on the CMB. No specific evidence prohibits a synchrotron mechanism, but such emission would imply much more severe energetic and lifetime constraints, while the IC/CMB process implies that the source will appear to have the same surface brightness at any redshift (Schwartz, 2002). The ratio of synchrotron (radio) emission to IC (X-ray) emission from a population of electrons (Jones, 1965; Felten & Morrison, 1966) is

$$\frac{L_R}{L_X}=\frac{\mathrm{H}^2/(8\pi )}{\rho _{CMB}}.$$
(2)
Since the energy density of microwave photons at the quasar is $`\rho _{CMB}`$ = aT4 = 7.56$`\times `$10-15 T$`{}_{0}{}^{4}(1+z)_{}^{4}`$
=10-9ergs cm-3 for the microwave temperature of T0 = 2.728 K (Fixsen et al., 1996) and redshift z=5.99, $`\rho _{CMB}`$ will dominate any magnetic field of strength less than 160$`\mu `$G. But from the radio flux upper limit, we can derive a rest frame 100 MHz to 100 GHz radio luminosity of less than 3$`\times `$1044 ergs s-1, compared to the rest frame 3.5–49 keV X-ray luminosity of 1.3$`\times `$1045 ergs s-1, and deduce that H$``$80$`\mu `$G from equation 2.

Electrons with $`\gamma `$ 1000 will broadly scatter the CMB photons to observed energies around 1 keV. Their lifetime will be 2.1$`\times `$1012/($`\gamma `$ (1+z)4) years. Taking the volume corresponding to a cylinder 5$`\mathrm{}`$ long and 1$`\mathrm{}`$ diameter at the source distance, 1.1$`\times `$1068cm3, gives a required energy density of 3.2$`\times `$10-10 ergs cm-3 in relativistic electrons emitting the X-rays. A 90 $`\mu `$G field would be in equipartition with such an electron density. The magnetic field could be a factor of a few smaller, due to uncertainties in the quantities or the modeling assumptions, and since there is not yet a measured radio flux.

We would typically expect fields of only tens of $`\mu `$G based on observations of X-ray jets to date (Harris & Krawczynski, 2002; Sambruna et al., 2002). Those observations have often required that the jet be moving with a bulk Lorentz factor $`\mathrm{\Gamma }`$ of at least a few, (see Harris & Krawczynski (2002); Sambruna et al. (2002)), in order that the CMB energy density be enhanced in the rest frame of the jet by a factor $`\mathrm{\Gamma }^2`$, and still preserve equipartition of the magnetic field and relativistic particle energy densities. Because of the large redshift we need not invoke relativistic beaming to explain the present observations. However, it is certainly allowed, as long as the jet is pointed within an angle $`\mathrm{arccos}(\sqrt{(}\frac{\mathrm{\Gamma }1}{\mathrm{\Gamma }+1}))`$ of our line of sight, so that the apparent flux is not diminished.

That the jet flux is about 40% of the quasar core flux in this case is consistent with the ratios of roughly 1% commonly observed in X-ray jets222But note that we so far do not have extensive, systematic X-ray surveys of jet properties. if we assume that the jet magnetic field is approximately 30 $`\mu `$G. In that case, if this object were at a redshift z=2, the core flux would be enhanced a factor of 14. The jet surface brightness would remain the same due to compensating factors of (1+z)4 in the microwave energy density and the cosmological dimming, and the solid angle would be about a factor of 2 smaller, for a net decrease in the jet to core ratio to about 1.5%. At redshifts less than 2, if it were not relativistically beamed, the CMB energy density would be less than the magnetic field energy density and the intrinsic X-ray emissivity of the jet would no longer be affected by the CMB.

Although the putative jet is about 30 kpc long, projected on the sky, there is an apparent gap of 130 kpc from the core to the jet, where the surface brightness is a factor of at least two smaller. This could be explained if the energy flux in this portion of the jet is primarily in the form of protons or Poynting flux, (cf. Harris & Krawczynski (2002)), or if it is mildly relativistic but away from our line of sight. In the latter case, the portion we see could be isotropic radiation after the jet becomes sub-relativistic.

## 4 CONCLUSIONS

All three z$``$6 quasars were detected in relatively short *Chandra* observations. Their X-ray to optical luminosity ratios are median values for high redshift, high luminosity quasars, indicating that quasars at higher Lx/LO, which are known to exist, will be detectable at if they exist at larger redshifts. Followup observations of all will be important to obtain rough spectral information, to investigate if SDSS 1030+0524 is gravitationally lensed, and to look more deeply for X-ray jets. In particular, the candidate jet from SDSS 1306 could be confirmed, or contradicted, by the spatial structure of 100 photons which would obtain in a 100 Ks observation. Radio emission should be detectable at $``$100$`\mu `$Jy at 5 GHz, by scaling from PKS 0637–752. This could be detected in a few hour VLA BnA observation. If confirmed, this will clearly indicate the possibility of detecting *only* the jet, and not core, X-ray emission from similar quasars at even larger redshifts.

This work was supported by NASA contract NAS8-39073 to the *Chandra* X-ray Center. I thank W. Forman, C. Jones, A. Siemiginowska, H. Tananbaum, and B. Wilkes for discussion and comments on the manuscript. A. Loeb pointed out the possibility that SDSS 1030 could be a gravitationally lensed system. This research used the NASA Astrophysics Data System Bibliographic Services, and the NASA/IPAC Extragalactic Database (NED) which is operated by the Jet Propulsion Laboratory, California Institute of Technology, under contract with the National Aeronautics and Space Administration.


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# Two-Component Fluid Membranes Near Repulsive Walls: Linearized Hydrodynamics of Equilibrium and Non-equilibrium States

## I Introduction

Amphiphilic molecules, in polar solvents such as water, form symmetric bilayer membrane phases at sufficient concentration . Such membranes are typically fluid in nature, with short-ranged positional order, although more complex forms of ordering are possible. The equilibrium conformations of such single-component membranes are governed by the energy cost for bending the bilayer, provided the surface tension can be neglected, as is the case if the membrane is self-assembled . Our understanding of the static and dynamic properties of such single-component membrane systems in equilibrium is now fairly detailed .

More complex multi-component membranes have also been synthesized and studied . Such membranes are especially important in the biological context, since cell membranes are often usefully idealized as a bilayer fluid-mosaic comprised of over a hundred different types of lipid and protein constituents. Biological membranes, however, often have additional complicating features: An underlying network of cross-linked proteins anchored to the bilayer and associated with the cell cytoskeleton typically lends such membranes a non-vanishing (although small) shear modulus . Another aspect, specific to the biological context, is the presence of non-equilibrium driving forces: An important class of trans-membrane proteins are the “ion pumps”, molecules which consume energy derived from ATP hydrolysis, electro-chemical gradients or light, and undergo conformational changes. Such pumps maintain electro-osmotic potential gradients across the cell membrane by controlling the flow of ions such as $`K^+`$ and $`Na^+`$. Since energy must be supplied externally, the driving of the pump is a process which occurs out of thermal equilibrium. Thus, the situation of active pumps diffusing in a fluid membrane matrix is an intrinsically non-equilibrium problem whose behavior represents a novel class of non-equilibrium steady states.

The appropriate statistical description of active or “driven” biological membrane systems has attracted recent experimental and theoretical attention. It has been realized that non-equilibrium behavior may underlie aspects of bio-membrane dynamics  previously attributed purely to equilibrium thermal fluctuations, such as the “flicker” phenomenon in erythrocytes. Recent micropipette experiments on light-activated bacteriorhodopsin (BR) pumps incorporated in phospholipid (EPC) bilayers find that area fluctuations in such active membrane systems can be phenomenologically described in terms of an effective temperature which exceeds the true physical temperature by up to a factor of 2 . In these systems, the transition between active and inactive states can be easily (and reversibly) tuned, providing a remarkable window into the dynamics of steady states away from equilibrium .

Analytical calculations which deal with the complex nature of a typical biological membrane in toto are difficult, if not impossible. We may, however, hope to gain useful intuition by working with simpler models. We idealize a typical biological membrane here as composed of principally two different types of molecular constituents, the lipids which constitute the bulk of the bilayer membrane and proteins which diffuse freely on the membrane surface. Such simple lipid/pump protein systems can be reconstituted and studied in vitro, as in the micropipette experiments referred to above. Our results should apply, most directly, to such experiments.

The question we address in this paper is the following: How are the fluctuations of such a two-component membrane, both in equilibrium and out of equilibrium, affected by the presence of a nearby repulsive wall? Fluctuations of a single-component membrane bounded on one side by a repulsive wall have been considered in earlier work . More recently, a similar problem for active membranes has been studied in an influential paper by Prost, Manneville and Bruinsma , who suggest that steady-state fluctuations of non-equilibrium membranes near repulsive walls can be amplified substantially relative to the equilibrium case. This amplification is studied via a scaling treatment which incorporates non-linear effects. Our starting model differs from theirs in several respects, in particular in the way we describe activity, in our incorporation of curvature-concentration coupling, as well as in our evaluation of the relative importance of permeative and hydrodynamic effects at the length scales accessed by typical experimental probes. We study the linearized hydrodynamics of such active pump-membrane systems in this paper, ignoring the role of non-linear effects. We will describe the similarities as well as the differences in the results we obtain.

We answer the question posed in the previous paragraph first through a calculation of the correlations and linearized mode structure in thermal equilibrium of a model impermeable two-component fluid membrane, incorporating hydrodynamic interactions. This amounts, in terms of the model for biological membranes discussed above, to assuming that the pump proteins are “passive” constituents of the membrane. While motivated here in a biological context, such models are also useful outside this specific context. Physics similar to that described in the case where the pumps are rendered passive should be generic to two-component systems composed of (i) a “symmetric” (lipid) constituent forming an up-down symmetric bilayer phase in isolation, and (ii) an asymmetric (protein) constituent, in which either the shape of the molecule or its location with respect to the bilayer favors a local spontaneous curvature for the bilayer. For concreteness, we will often use the terms lipid and protein when referring to constituents of types (i) and (ii), following our earlier discussion, but stress that the results obtained here in the equilibrium case are also applicable more generally.

For protein molecules which lack up-down symmetry, we may distinguish between molecules oriented parallel to the local normal and those which are oriented antiparallel to the normal. We label these proteins as “+” and “-” for convenience; the relatively small flip-flop rate for protein transfer across leaves of the bilayer ensures that these labels are conserved across short and intermediate time scales. The difference between the local densities of + and - proteins defines a “signed ” protein density field $`\psi `$. To describe the incorporation of such asymmetric proteins into the lipid bilayer, we model the protein as a rigid rod of length $`w^{}+w^{}`$. The protein is taken to situate itself asymmetrically with respect to the bilayer midpoint; a section $`w^{}`$ of the protein lies on one side of the bilayer midpoint while a section $`w^{}`$ lies on the other side. This relatively simple model of the asymmetry is convenient for analytic computation.

To lowest order, the two-component character of the membrane manifests itself via the existence of a curvature-concentration coupling – fluctuations in $`\psi `$ influence the mean curvature in that region (see Fig. 1). We will work with balanced membranes for which $`\psi =0`$. For inactive pumps, corresponding to the equilibrium case, the curvature-concentration coupling term in the free energy effectively shifts the bending rigidity of the membrane from $`\kappa `$ to $`\kappa _e=\kappa (\kappa \overline{H})^2/\chi _{0}^{}{}_{}{}^{1}`$. Here $`\overline{H}`$ is the coefficient coupling curvature to concentration and $`\chi _{0}^{}{}_{}{}^{1}`$ an inverse compressibility .

In closed vesicular structures found in biological contexts, while the bilayer constituting the vesicle may be symmetric, chemical and physical environments within and outside the vesicle can differ considerably. Thus, a pump protein can, in principle, distinguish the side of the bilayer exposed to the inner volume from the side which is exposed to the outer volume. For a large enough vesicle, flat on the large length scales of relevance to a hydrodynamic calculation, the pump molecule can thus tell up from down, where up and down are defined arbitrarily with respect to the average position and orientation of the bilayer midpoint. As an extreme limit, we may consider the case in which the pump molecules preferentially insert always on the same (say up) side, so that a section $`w^{}`$ of the protein molecule always lies above the bilayer midpoint while a section $`w^{}`$ lies below.

A recent study of active membranes works with this model of the architecture of a membrane-pump system, modeling activity in terms of dipole forces associated with the ends of the pump, as we detail below . The associated forces may point either outward at the ends (up pumps) or inward at the ends (down pumps); see Fig. 2. In this model, up and down pumps are physically distinguishable. The main results presented here are appropriate to this model.

Note that this distinction between “above” and “below” is artificial for strictly symmetric bilayer membranes, in which pump proteins may insert on either leaf of the bilayer with equal probability. In an Appendix, we discuss results for an alternative model, in which the protein attachment respects the “up-down” symmetry of the bilayer; the center of mass of the protein can be displaced either above or below the bilayer mid-plane as shown in Fig. 3. The head and tail distances from the bilayer midpoint are fixed at $`w_l`$ and $`w_s`$ respectively. In this model, up pumps and down pumps are physically indistinguishable; their nomenclature follows from the side of the bilayer on which they are inserted.

We will present results for non-equilibrium membranes in the bulk, in which case the pumps are active, and for the case in which fluctuations of such an active membrane are bounded on one side by a repulsive wall. This study complements results by Prost, Manneville and Bruinsma for a model in which curvature-concentration coupling is ignored, an effect explicitly incorporated in later work . In the active case, the pumps exert forces on the membrane and the surrounding fluid. Since these forces are internal forces, they must cancel when integrated over the size of the pump. Following Manneville et al. in Ref., we consider the simplest model for pumps consistent with this requirement: the pumps are taken to be dipole force centres with positive and negative force centres located asymmetrically with respect to the midpoint of the bilayer.

Our other major assumptions are the following: We assume that our observations are conducted on scales such that the membrane may be considered to be impermeable to the solvent in which it is embedded. This assumption follows from the work of Manneville et al. who observe that for active terms permeative effects can be neglected vis. a vis hydrodynamic ones over a substantial range of length scales upto microscopic ones. We derive our results under the assumption that the active membrane is impermeable and extend this assumption to the case when the membrane is in the passive state, to smoothly interpolate between results in the two regimes. (In the Rouse or free draining limit, where the membrane dynamics decouples from that of the fluid, the membrane is implicitly permeable). We take the solvent to be incompressible and work at small Reynolds number, neglecting the inertia of the fluid. We use linearized equations of motion for the hydrodynamic velocity field, corresponding to the Stokes limit of the Navier-Stokes equations.

We now summarize our principal results. For equilibrium, impermeable, two-component membranes bounded on one side by a wall placed at a distance $`d`$ from the membrane plane, we obtain the following modified mode structure to leading order,

$$\omega _1𝒢(\sigma k_{}^2+\kappa _ek_{}^4),\omega _2Dk_{}^2.$$
(1)
In our notation, $`D`$ is the diffusion coefficient of the proteins on the membrane, $`\sigma `$ the surface tension of the membrane, $`\kappa `$ the bending rigidity of the membrane and $`\eta `$ the viscosity of the fluid. Here,

$`𝒢(k_{},k_{}d)`$ $``$ $`\{\begin{array}{cc}d^3k_{}^2/12\eta \hfill & \text{if }k_{}1/d,\hfill \\ 1/4\eta k_{}\hfill & \text{if }k_{}1/d.\hfill \end{array}`$ (4)
is a“crossover function”, which describes how membrane fluctuations interpolate between a regime where physical interactions between the membrane and the wall cannot be neglected, and a regime where such fluctuations do not feel the presence of the wall. In the $`k_{}1/d`$ limit, the above result reduces to one derivable in the bulk case, as is physically sensible.

To study non-equilibrium membranes we model active pumps, following Manneville et al., as “asymmetric” force dipoles with force-centres located at distances $`w^{}`$ above the bilayer midpoint and $`w^{}`$ below the bilayer midpoint. These distances are, in general, unequal. The magnitude of the force exerted at each of these force-centres is $`f`$. The pumps diffuse on the membrane and are characterized by a mobility $`\mathrm{\Lambda }`$.

In the presence of a wall, for non-equilibrium membranes, it has been argued that static height fluctuations are amplified and an interesting speculation is that such an amplification may, in fact, speed up the process of membrane fusion when two non-equilibrium membranes are brought close together . When the wall is at a distance $`d`$ from the membrane, our calculation yields (for a tensionless membrane)

$`\omega _1`$ $``$ $`{\displaystyle \frac{f𝒢^{aw}\kappa \overline{H}}{\chi _{0}^{}{}_{}{}^{1}}}k_{}^2+𝒢\kappa _ek_{}^4,`$ (5)
$`\omega _2`$ $``$ $`Dk_{}^2+{\displaystyle \frac{\kappa \overline{H}(\kappa \overline{H}𝒢k_{}^2f𝒢^{aw})}{\chi _{0}^{}{}_{}{}^{1}}}k_{}^2.`$ (6)
where $`𝒢^{aw}`$ is the crossover function

$`𝒢^{aw}(k_{},d,w^{},w^{})`$ $``$ $`\{\begin{array}{cc}dw_{}^{}{}_{}{}^{2}k_{}^2/4\eta \hfill & \text{if }k_{}1/d,\hfill \\ \mathrm{\Omega }(k_{},w^{},w^{})/4\eta k_{}\hfill & \text{if }k_{}1/d.\hfill \end{array}`$ (9)
$`\mathrm{\Omega }(k_{},w^{},w^{})`$ is proportional to the degree of asymmetry $`(w^{}w^{})`$. In a regime where fluctuations feel the wall, non-equilibrium effects do not depend on the asymmetry factor but only on $`w^{}`$, as seen above. Here again, curvature-concentration coupling plays a crucial role in the manifestation of non-equilibrium effects in the linearized mode structure.

We find that in the presence of the wall, the first term in $`\omega _1`$ is the most important since it is of order $`O(k_{}^4)`$. The existence of this mode depends crucially on the presence of both activity and curvature-concentration coupling. It will be shown later in this paper that this mode causes active membranes near walls to relax faster than their equilibrium counterparts and causes a smoothening of membrane fluctuations; in the presence of a wall, the roughness of tensionless, active membranes averaged over a patch of size $`L\times L`$, ($`h^2(L)`$), scales as ln(L) instead of $`L^2`$. Thus the wall acts to smoothen active membrane fluctuations in a tensionless membrane by inducing an apparent surface tension. This result is a purely non-equilibrium one, since the apparent surface tension vanishes when the strength of active forces is set to zero. It also relies crucially on the existence of a curvature-concentration coupling, since the apparent surface tension also vanishes when the curvature and concentration fields are decoupled. We show that this effect can be understood in terms of the dynamics of the fast relaxing $`\psi `$ field which drives the height field $`h(𝐫_{},t)`$ via activity and curvature-concentration coupling.

The outline of this paper is as follows: Section II describes our basic coarse-grained free energy and reviews the calculation of the static properties of a two-component membrane with curvature-concentration coupling. In Section III (a), the free-draining approximation (Rouse model) is used to discuss the dynamics. In Section III (b), we incorporate the effects of the hydrodynamic velocity field of the solvent. Section IV describes the mode structure in the presence of a wall for the equilibrium problem. In Section V, we discuss the mode structure and correlation functions of active two-component membranes. Section VI discusses how these properties are modified by the presence of a wall. In Section VII, we summarize the conclusions of this paper. An Appendix discusses the details of an alternative model for active two-component membranes, with up-down symmetry.

## II Statics

We model the membrane as an infinitesimally thin two-dimensional surface embedded in three-dimensional space. Points on this surface are specified by a three-dimensional vector $`𝐑(\stackrel{~}{u})`$, with components $`R_i(\stackrel{~}{u}),i=1\mathrm{}3`$. We use Monge gauge, valid for nearly flat surfaces, in which the surface is specified by its height above a two-dimensional flat plane. This plane is parameterized via Euclidean coordinates i.e. $`\stackrel{~}{u}=(x,y)𝐫_{}`$; $`𝐑(𝐫_{})=(x,y,h(x,y))`$. In this gauge, the curvature is $`H=^2h`$, to lowest order.

The number densities of the lipid molecules, + proteins and - proteins are specified through fields $`\rho _{\mathrm{}}`$, $`\rho ^+(\stackrel{~}{u},t)`$ and $`\rho ^{}(\stackrel{~}{u},t)`$ respectively. These are the physical densities of the lipids and proteins on the membrane. The projections of these densities on the $`xy`$ plane are obtained by multiplying the physical density fields by the metric factor $`\sqrt{1+(h)^2}`$. To lowest order in gradients of $`h`$, physical densities and projected densities coincide. The scalar fields $`\varphi (\stackrel{~}{u},t)`$ and $`\psi (\stackrel{~}{u},t)`$ are defined by $`\varphi (\stackrel{~}{u},t)=\rho ^+(\stackrel{~}{u},t)+\rho ^{}(\stackrel{~}{u},t)`$ and $`\psi (\stackrel{~}{u},t)=\rho ^+(\stackrel{~}{u},t)\rho ^{}(\stackrel{~}{u},t)`$.

The free energy for the membrane-protein system consists of a part arising from protein and lipid densities (the matter part) and another which arises as a consequence of membrane fluctuations about the flat state. Given equilibrium concentrations of lipid and protein ($`\rho _0`$), $`\varphi (\stackrel{~}{u},t)=\rho ^+(\stackrel{~}{u},t)+\rho ^{}(\stackrel{~}{u},t)=\varphi _0`$ , $`\psi (\stackrel{~}{u},t)=\rho ^+(\stackrel{~}{u},t)\rho ^{}(\stackrel{~}{u},t)=\psi _0`$ and fixing $`\rho \rho _0`$ and $`\varphi \varphi _0`$, the appropriate gauge-invariant free energy of the membrane with asymmetric proteins is

$`F`$ $`=`$ $`{\displaystyle d^2\stackrel{~}{u}\sqrt{g}[\sigma +\frac{1}{2}\chi _0^1(\psi \psi _0)^2]}+{\displaystyle \frac{1}{2}}\kappa {\displaystyle d^2\stackrel{~}{u}\sqrt{g}(HH_0)^2}.`$ (10)
where $`\sigma `$ is the surface tension and $`\kappa `$ is the bending rigidity. To lowest order we take $`H_0=\overline{H}\psi `$. To leading order in gradients of $`h`$, we have

$$F=\frac{1}{2}d^2𝐫_{}[\kappa (^2h(𝐫_{},t))^2+\sigma (h(𝐫_{},t))^22\overline{H}\kappa \psi (𝐫_{},t)^2h(𝐫_{},t)+\chi _0^1\psi (𝐫_{},t)^2],$$
(11)
yielding a local force density of $`𝐟(𝐫_{})=\delta F/\delta h|_\psi `$. The local force density is evaluated at constant density .

The partition function is

$$Z=𝒟[h]𝒟[\psi ]e^{\beta F[h,\psi ]},$$
(12)
which yields, on integrating out the $`\psi `$ field, an effective free energy for the height field $`F_{eff}[h]`$,

$$F_{eff}[h]=\frac{1}{2}\underset{k_{}}{}(\sigma k_{}^2+\kappa k_{}^4)|h(k_{})|^2\frac{(\kappa \overline{H})^2}{\chi _{0}^{}{}_{}{}^{1}}k_{}^4|h(k_{})|^2.$$
(13)
The equipartition theorem yields the equilibrium correlation function for the height field fluctuations (about the equilibrium flat, “mixed” phase) as

$$h^{}(𝐤_{})h(𝐤_{})=\frac{k_BT}{(\sigma k_{}^2+\kappa _ek_{}^4)},$$
(14)
where $`\kappa _e=\kappa (\kappa \overline{H})^2/\chi _{0}^{}{}_{}{}^{1}`$. The asymmetric components act to reduce the rigidity modulus from $`\kappa `$ to $`\kappa _e`$ as a consequence of curvature-concentration coupling , an effect easily understood on physical grounds. We assume that $`\overline{H}`$ is small enough to keep $`\kappa _e`$ positive, in which case the membrane is linearly stable. Instabilities arising at equilibrium due to large values of $`\overline{H}`$ have been discussed in .

## III Mode structure in the bulk

We now study the dynamics of a two-component membrane with curvature-concentration coupling and suspended in a solvent. We will first present results in the Rouse model limit in which the background fluid drains freely through the membrane and effects due to the hydrodynamics of the background fluid velocity field can be ignored .

### A Rouse Model

In this model the fluid velocity field decouples from the membrane height field and $`\psi `$. The membrane is implicitly permeable and the height field fluctuations are governed by a balance between the elastic forces and local (permeative) friction. An equation of the form

$$_th(𝐫_{},t)=\mathrm{\Gamma }\frac{\delta F}{\delta h}+f_m,$$
(15)
follows, where $`\mathrm{\Gamma }`$ is a kinetic coefficient and $`f_m`$ is a thermal equilibrium noise whose statistics ensures that the static height-height correlation function is Eq. (14).

The proteins diffuse freely on the surface of the membrane, Since $`\psi `$ is locally conserved, it obeys an equation of continuity of the form

$$_t\psi (𝐫_{},t)=\mathrm{\Lambda }^2\frac{\delta F}{\delta \psi }+𝐟_\psi ,$$
(16)
with $`\mathrm{\Lambda }=D/\chi _{0}^{}{}_{}{}^{1}`$, D is the diffusion coefficient of the proteins on the membrane. A contribution due to the in-plane current of the pumps has been dropped; this is typically small and can be neglected to linear order. This term contributes if we go beyond the strictly linear treatment by replacing $`\psi ^2`$ by $`\psi ^2`$, yielding a convective term which gives rise to waves whose speed is independent of the wavevector . The last term is a conserving Gaussian noise with $`𝐟_\psi (𝐫_{},𝐭)=\mathrm{𝟎}`$ and correlations $`f_{\psi i}(𝐫_{},t)f_{\psi j}(𝐫_{}^{},t^{})=2\mathrm{\Lambda }k_BT\delta _{ij}\delta (𝐫_{}𝐫_{}^{})\delta (tt^{})`$.

Equations (15) and (16) can be written as

$`_t\left(\begin{array}{c}h(𝐤_{},\omega )\\ \psi (𝐤_{},\omega )\end{array}\right)`$ $`=`$ $`\left(\begin{array}{cc}\mathrm{\Gamma }(\sigma k_{}^2+\kappa k_{}^4)& \mathrm{\Gamma }\kappa \overline{H}k_{}^2\\ \mathrm{\Lambda }\kappa \overline{H}k_{}^4& \mathrm{\Lambda }\chi _0^1k_{}^2\end{array}\right)\left(\begin{array}{c}h(𝐤_{},\omega )\\ \psi (𝐤_{},\omega )\end{array}\right)+\left(\begin{array}{c}f_m\\ i𝐤_{}.𝐟_\psi \end{array}\right).`$ (25)
Solving these coupled equations yields

$`h^{}(𝐤_{},\omega )h(𝐤_{},\omega )={\displaystyle \frac{f_mf_m(\omega ^2+\tau _{\psi }^{1}{}_{}{}^{2})+2(\mathrm{\Gamma }\kappa \overline{H})^2\mathrm{\Lambda }k_BTk_{}^6}{(\omega ^2+\tau _h^1\tau _\psi ^1\mathrm{\Lambda }\mathrm{\Gamma }(\kappa \overline{H})^2k_{}^6)^2+\omega ^2(\tau _\psi ^1+\tau _h^1)^2}},`$ (26)
where $`\tau _h^1=\mathrm{\Gamma }(\sigma k_{}^2+\kappa k_{}^4)`$ and $`\tau _\psi ^1=\mathrm{\Lambda }\chi _0^1k_{}^2`$.

On integrating over $`\omega `$, this correlation function should reduce to the same form as Eq. (14). This constrains the equilibrium thermal noise correlations to be the following:

$$f_m(𝐫_{},t)f_m(𝐫_{}^{},t^{})=2\mathrm{\Gamma }k_BT\delta (𝐫_{}𝐫_{}^{})\delta (tt^{}).$$
(27)
In addition, we impose $`f_m(𝐫_{},t)=0`$.

To obtain the mode structure, we drop the noise terms in Eq. (25) and find the eigenvalues of the matrix. This yields

$`\omega _1,\omega _2`$ $`=`$ $`{\displaystyle \frac{1}{2}}\left(\mathrm{\Gamma }(\sigma k_{}^2+\kappa k_{}^4)+Dk_{}^2\right)\pm {\displaystyle \frac{1}{2}}\left(\sqrt{(\mathrm{\Gamma }(\sigma k_{}^2+\kappa k_{}^4)+Dk_{}^2)^24D\mathrm{\Gamma }k_{}^2(\sigma k_{}^2+\kappa _ek_{}^4)}\right)`$ (28)
In the absence of curvature-concentration coupling these modes decouple and are $`\omega _1,\omega _2`$ = $`\mathrm{\Gamma }(\sigma k_{}^2+\kappa k_{}^4)`$, $`Dk_{}^2`$, the dispersive modes of a tense membrane . For tensionless membranes, the presence of curvature-concentration coupling modifies the mode structure to

$$\omega _1\mathrm{\Gamma }\kappa _ek_{}^4,\omega _2Dk_{}^2+\frac{(\kappa \overline{H})^2}{\chi _{0}^{}{}_{}{}^{1}}\mathrm{\Gamma }k_{}^4.$$
(29)
### B Zimm Model

Next, we examine the effect of solvent hydrodynamics on the dynamical correlation functions of the protein density field and the membrane height field . Since no external forces act on the (fluid+membrane+proteins) system, momentum conservation requires

$$_tg_i=_j\pi _{ij},$$
(30)
where $`𝐠`$ is the momentum density and $`\pi _{ij}`$ is the stress tensor. The stress tensor takes the form $`\pi _{ij}=P\delta _{ij}+\rho v_iv_j\eta (_iv_j+_jv_i)+𝒮_{ij},`$ where $`𝐯(𝐫,t)`$ is the fluid velocity, $`P(𝐫,t)`$ is the pressure field and $`𝒮_{ij}`$ describes the contribution to the stress tensor arising from membrane conformations (the $`^2`$ in the above equation is evaluated in 3 dimensions). In the Stokes limit, the non-linear term can be dropped. At low Reynolds number, we neglect the inertial term on the left hand side of Eq. (30) leading to $`_j\pi _{ij}=0`$, and thus the equation governing the hydrodynamic velocity field

$$\eta ^2𝐯(𝐫,t)P(𝐫,t)𝐅(𝐫,t)+𝐟_𝐡(𝐫,t)=0.$$
(31)
$`𝐅(𝐫,t)`$ is the force acting on the fluid due to membrane stresses $`𝒮_{ij}`$ and $`𝐟_𝐡(𝐫,t)`$ is the equilibrium thermal noise present in the bulk of the fluid. This noise obeys $`𝐟_𝐡(𝐫,t)=0`$  and is related to the viscosity via the fluctuation-dissipation theorem.

$$f_{hi}(𝐫,t)f_{hj}(𝐫^{},t^{})=2k_BT\eta \{\delta _{ij}^2+_i_j\}\delta (𝐫𝐫^{})\delta (tt^{}),$$
(32)
where $`(i,j)`$ index the components of the force. Eqns. (31) are to be solved together with the incompressibility condition

$$𝐯(𝐫,t)=0,$$
(33)
to obtain the mode structure of an impermeable two-component membrane in the presence of an incompressible solvent.

We employ a useful convention due to Seifert in which a basis spanned by the $`z`$-component and the longitudinal and transverse in-plane components of the velocity is used . (This convention will be useful further on in this paper when we discuss membrane fluctuations in the presence of a confining wall). We solve the above equations to obtain the transverse ($`t`$), longitudinal ($`l`$) and $`z`$ components of the fluid velocity in terms of the $`t`$,$`l`$ and $`z`$ components of the force $`𝐅`$ acting on the fluid . The $`t`$ and $`l`$ components of the velocity lie in the plane of the membrane and are perpendicular and parallel to $`𝐤_{}`$, the wavevector describing fluctuations of the membrane. Some algebra yields these components as

$`v_z(𝐤_{},z)`$ $`=`$ $`{\displaystyle \frac{1}{4\eta k_{}}}{\displaystyle _{\mathrm{}}^{\mathrm{}}}𝑑z^{}e^{k_{}|z^{}z|}[(1+k_{}|zz^{}|)F_z(𝐤_{},z^{})+ik_{}(z^{}z)F_l(𝐤_{},z^{})],`$ (34)
$`v_l(𝐤_{},z)`$ $`=`$ $`{\displaystyle \frac{1}{4\eta k_{}}}{\displaystyle _{\mathrm{}}^{\mathrm{}}}𝑑z^{}e^{k_{}|z^{}z|}[(1k_{}|zz^{}|)F_l(𝐤_{},z^{})+ik_{}(z^{}z)F_z(𝐤_{},z^{})]`$ (35)
$`v_t(𝐤_{},z)`$ $`=`$ $`{\displaystyle \frac{1}{4\eta k_{}}}{\displaystyle _{\mathrm{}}^{\mathrm{}}}𝑑z^{}e^{k_{}|z^{}z|}2F_t(𝐤_{},z^{}).`$ (36)
Note that the transverse components can be ignored since they do not couple to the other components. We can now calculate $`v_z(𝐤_{},0)`$ by supplementing the above equations with boundary conditions.

In the absence of permeation the membrane is advected by the fluid. The velocity of any point on the membrane is the $`z`$-component of the velocity of the fluid at that point. The equation of motion for the height field of an impermeable membrane is therefore

$$_th(𝐫_{},t)=v_z(𝐫_{},0,t),$$
(37)
where $`v_z(𝐫_{},0,t)`$ is the velocity of the surrounding fluid at the mean position of the membrane. The force acting on the fluid is of the form

$$𝐅(𝐤_{},z)=\delta (z)[\frac{\delta F}{\delta h}\widehat{𝐳}+F_l^m\widehat{𝐤_{}}],$$
(38)
where $`F_l^m`$ is the longitudinal component of the force acting on the fluid due to the membrane. The elastic force density due to the membrane is $`\delta F/\delta h\widehat{𝐳}`$ . We solve for $`v_z(𝐤_{},0)`$ using Eqns. (34), together with the boundary condition that the membrane is incompressible i.e. the in-plane divergence of the velocity field vanishes at the membrane. This implies that $`v_l(𝐤_{},0)=0`$. This boundary condition gives $`F_l^m=0`$, yielding

$$v_z(𝐤_{},0)=\frac{1}{4\eta k_{}}\frac{\delta F}{\delta h}.$$
(39)
The bulk thermal noise present in the fluid also contributes to $`v_z(𝐤_{},0)`$. This contribution is denoted by $`f_z`$, with

$$f_z(𝐤_{},0)=\frac{dk_z}{2\pi \eta k^2}(\delta _{jz}\frac{k_jk_z}{k^2})[f_h(𝐤,t)]_j.$$
(40)
We can use the noise correlation function given in Eq. (32) to show that the noise $`f_z`$ has zero mean and that its variance in Fourier space is $`2k_BT/4\eta k_{}`$.

Finally, Eq. (37) becomes

$$_th=\frac{1}{4\eta k_{}}\frac{\delta F}{\delta h}+f_z(𝐤_{},0,t).$$
(41)
The density difference field obeys Eq. (16). Eq. (37) and Eq. (16) can then be written as

$`_t\left(\begin{array}{c}h(𝐤_{},\omega )\\ \psi (𝐤_{},\omega )\end{array}\right)`$ $`=`$ $`\left(\begin{array}{cc}\frac{\sigma k_{}^2+\kappa k_{}^4}{4\eta k_{}}& \frac{\kappa \overline{H}k_{}^2}{4\eta k_{}}\\ \mathrm{\Lambda }\kappa \overline{H}k_{}^4& \mathrm{\Lambda }\chi _0^1k_{}^2\end{array}\right)\left(\begin{array}{c}h(𝐤_{},\omega )\\ \psi (𝐤_{},\omega )\end{array}\right)+\left(\begin{array}{c}f_z(𝐤_{},\omega )\\ i𝐤_{}.𝐟_\psi \end{array}\right).`$ (50)
Solving these equations gives

$`h^{}(𝐤_{},\omega )h(𝐤_{},\omega )={\displaystyle \frac{(2k_BT/4\eta k_{})(\omega ^2+\tau _{\psi }^{1}{}_{}{}^{2})+(k_BT\mathrm{\Lambda }(\kappa \overline{H})^2/8\eta ^2)k_{}^4}{(\omega ^2+\tau _h^1\tau _\psi ^1(\mathrm{\Lambda }(\kappa \overline{H})^2/4\eta )k_{}^5)^2+\omega ^2(\tau _\psi ^1+\tau _h^1)^2}}.`$ (51)
The parameters appearing in the above equations are

$$\tau _h^1=\frac{1}{4\eta k_{}}(\sigma k_{}^2+\kappa k_{}^4),\tau _\psi ^1=\mathrm{\Lambda }\chi _0^1k_{}^2.$$
(52)
Integrating this result over $`\omega `$ recovers for us the equal time correlation function for height fluctuations about the equilibrium, flat, “mixed” phase

$$h^{}(𝐤_{},t)h(𝐤_{},t)=\frac{k_BT}{(\sigma k_{}^2+\kappa _ek_{}^4)},$$
(53)
as expected. The roughness of the height field at macroscopic length scales $`L`$, defined through $`h^2(L)=_{1/L}^{1/a}d^2k_{}h(k_{})h^{}(k_{})`$ scales as

$`h^2(L)=\{\begin{array}{cc}\frac{k_BT}{\kappa _e}L^2\hfill & \text{if }\sigma =0,\hfill \\ \frac{k_BT}{\sigma }ln(L/a)\hfill & \text{if }\sigma 0,\hfill \end{array}`$ (56)
where a is a microscopic cut-off length.

The mode structure is then

$`\omega _1,\omega _2`$ $`=`$ $`{\displaystyle \frac{1}{8\eta }}\left(\sigma k_{}+\kappa k_{}^3+4\eta Dk_{}^2\right)\pm {\displaystyle \frac{1}{8\eta }}\left(\sqrt{(\sigma k_{}+\kappa k_{}^3+4\eta Dk_{}^2)^216\eta D(\kappa _ek_{}^5+\sigma k_{}^3)}\right).`$ (57)
In the absence of curvature-concentration coupling, $`\omega _1,\omega _2=(\sigma k_{}+\kappa k_{}^3)/4\eta ,Dk_{}^2`$, i.e. the decoupled modes of the membrane height field and the protein concentration field in the presence of hydrodynamic correlations. For tensionless membranes with non-zero curvature-concentration coupling, the modes are

$$\omega _1\frac{\kappa _e}{4\eta }k_{}^3,\omega _2Dk_{}^2+\frac{(\kappa \overline{H})^2}{4\eta \chi _{0}^{}{}_{}{}^{1}}k_{}^3.$$
(58)
## IV Mode Structure in the presence of a wall

Consider a fluid bilayer membrane with shape asymmetric constituents, suspended in a fluid bounded on one side by a wall. Our model for the membrane-pump system follows that of Manneville et al.. In our co-ordinates, the membrane fluctuates about the $`z=d`$ plane and the wall is at $`z=0`$. The membrane is kept at distance $`d`$ from the wall by a repulsive interaction with the wall and a suitable external pressure which prevents it from floating away into the bulk.

The relaxation rate of an equilibrium membrane is reduced near a wall; in tandem, the equilibrium noise fluctuations are required to have a reduced amplitude as a consequence of the fluctuation-dissipation theorem . On short-length scales, however, membrane fluctuations do not see the wall and the height-height correlation function is identical to that in the bulk. This can be used to determine the form of thermal correlation functions in the presence of the wall.

We work in the linear regime where $`h^2(L_c)d^2`$, defining a collision length $`L_c`$. Beyond this length the membrane senses the presence of the wall and the non-linearities neglected in our treatment become important; $`1/L_c`$ is the smallest wave vector above which the linearization condition $`|h(𝐫_{},t)|d`$ is valid. From Eq. (56) we can estimate the collision length of an equilibrium membrane near a wall to be

$`L_c(d)=\{\begin{array}{cc}\sqrt{\frac{\kappa _e}{k_BT}}d\hfill & \text{if }\sigma =0,\hfill \\ aexp(\sigma d^2/k_BT)\hfill & \text{if }\sigma 0.\hfill \end{array}`$ (61)
We now solve Eq. (31) to obtain the velocity of the fluid at the position of the membrane. The membrane and wall are introduced in Eq. (31) as external forces acting on the fluid, imposing the required boundary conditions. The incompressibility of the fluid is used to solve for the components of the velocity in terms of the components of the external force.

We express the components of the velocity in the “mixed” basis of Seifert  as before. The force exerted by the membrane and wall on the fluid is of the form

$$𝐅(𝐤_{},z)=\delta (zd)[\frac{\delta F}{\delta h}\widehat{𝐳}+F_l^m\widehat{𝐤_{}}]+\delta (z)[F_z^w\widehat{𝐳}+F_l^w\widehat{𝐤_{}}],$$
(62)
where $`F_z^w`$ and $`F_l^w`$ are the $`z`$ and longitudinal components of the force exerted by the wall on the fluid at $`z=0`$. The quantities $`F_l^m`$, $`F_z^w`$ and $`F_l^w`$ can be evaluated using the boundary conditions: all components of the velocity should vanish at the wall (no-slip boundary condition) . This imposes

$$v_z(𝐤_{},0,t)=v_l(𝐤_{},0,t)=v_t(𝐤_{},0,t)=0.$$
(63)
These conditions, together with membrane incompressibility ($`v_l(𝐤_{},d,t)=0`$), provide the boundary conditions for the velocity field of the fluid at the membrane. This gives $`v_z(k_{},d,t)=𝒢\frac{\delta F}{\delta h}`$. The equation of motion for the height field is then obtained from Eqn. (37),

$$_th(𝐤_{},t)=𝒢\frac{\delta F}{\delta h}+\stackrel{~}{f_z}(𝐤_{},d,t),$$
(64)
where

$`𝒢(k_{},k_{}d)`$ $`=`$ $`{\displaystyle \frac{1}{4\eta k_{}}}\left({\displaystyle \frac{2(\mathrm{sinh}^2(k_{}d)(k_{}d)^2)}{\mathrm{sinh}^2(k_{}d)(k_{}d)^2+k_{}d+\mathrm{sinh}(k_{}d)\mathrm{cosh}(k_{}d)}}\right),`$ (65)
$``$ $`\{\begin{array}{cc}d^3k_{}^2/12\eta \hfill & \text{if }k_{}1/d{}_{}{}^{},\hfill \\ 1/4\eta k_{}\hfill & \text{if }k_{}1/d.\hfill \end{array}`$ (68)
We note that the term marked $``$ is incorrect for arbitrarily small wave vectors. The relevant cut-off is the inverse of the collision length $`L_c`$ since linearized calculations are not valid at wave-vectors smaller than this value.

The contribution to the velocity field at the membrane due to the noise in the fluid in the presence of the wall is $`\stackrel{~}{f_z}`$. Following the procedure used in the derivation of Eq. (50), we get

$`_t\left(\begin{array}{c}h(𝐤_{},\omega )\\ \psi (𝐤_{},\omega )\end{array}\right)=\left(\begin{array}{cc}𝒢(\sigma k_{}^2+\kappa k_{}^4)& 𝒢(\kappa \overline{H}k_{}^2)\\ \mathrm{\Lambda }\kappa \overline{H}k_{}^4& \mathrm{\Lambda }\chi _0^1k_{}^2\end{array}\right)\left(\begin{array}{c}h(𝐤_{},\omega )\\ \psi (𝐤_{},\omega )\end{array}\right)+\left(\begin{array}{c}\stackrel{~}{f_z}(𝐤_{},\omega )\\ i𝐤_{}.𝐟_\psi \end{array}\right).`$ (77)
This matrix equation is solved to yield

$`h^{}(𝐤_{},\omega )h(𝐤_{},\omega )={\displaystyle \frac{\stackrel{~}{f_z}\stackrel{~}{f_z}(\omega ^2+\tau _{\psi }^{1}{}_{}{}^{2})+2k_BT\mathrm{\Lambda }(𝒢\kappa \overline{H})^2k_{}^6}{(\omega ^2+\tau _{h}^{1}{}_{}{}^{w}\tau _\psi ^1𝒢\mathrm{\Lambda }(\kappa \overline{H})^2k_{}^6)^2+\omega ^2(\tau _\psi ^1+\tau _{h}^{1}{}_{}{}^{w})^2}},`$ (78)
where $`\tau _{h}^{1}{}_{}{}^{w}=𝒢(\sigma k_{}^2+\kappa k_{}^4)`$.

The hydrodynamics of the solvent cannot change equilibrium correlation functions. Thus, demanding that equal time correlations of the $`h`$ field are the same as in Eq. (53), we obtain the noise correlation function of $`\stackrel{~}{f_z}`$ as

$`\stackrel{~}{f_z}(𝐤_{},t)`$ $`=`$ $`0,`$ (79)
$`\stackrel{~}{f_z}(𝐤_{},t)\stackrel{~}{f_z}(𝐤_{}^{},t^{})`$ $`=`$ $`2k_BT𝒢(k_{},k_{}d)\delta (𝐤_{}+𝐤_{}^{})\delta (tt^{}).`$ (80)
Note that this reduces to the noise correlation function for the equilibrium bulk noise when $`k_{}>>1/d`$, i.e. when the membrane fluctuations are not significantly affected by the wall (see Section III).

We note here that the equation of motion of the height field can be derived in the limit $`1/L_c<<k_{}<<1/d`$ by directly imposing impermeability as in . Since the membrane is impermeable, the height field is conserved and obeys a continuity equation $`_th=_{}.j_h`$. The height “current” can be calculated from the lubrication approximation as $`j_h=\frac{d^3}{12\eta }_{}P`$, where $`P`$ is the pressure due to the elastic forces and the noise. Using the appropriate expressions for these terms, we recover the equation for the height (Eq. (64)) in the “close to the wall” limit.

The eigen-modes follow from Eq. (77)

$`\omega _1,\omega _2={\displaystyle \frac{1}{2}}\left(Dk_{}^2+\sigma 𝒢k_{}^2\pm \kappa 𝒢k_{}^4\right)+{\displaystyle \frac{1}{2}}\left(\sqrt{(\sigma 𝒢k_{}^2+Dk_{}^2+\kappa 𝒢k_{}^4)^24𝒢D(\sigma +\kappa _ek_{}^2)k_{}^4}\right)`$ (81)
In the absence of curvature-concentration coupling the modes corresponding to the hydrodynamic relaxation of a two-component membrane near a wall are recovered : $`\omega _1,\omega _2=𝒢(\sigma k_{}^2+\kappa k_{}^4),Dk_{}^2`$. For membranes with curvature-concentration coupling the modes are modified to

$$\omega _1𝒢(\sigma k_{}^2+\kappa _ek_{}^4),\omega _2Dk_{}^2+𝒢\frac{(\kappa \overline{H})^2}{\chi _{0}^{}{}_{}{}^{1}}k_{}^4.$$
(82)
It is easy to see that these reduce to the expressions derived in the bulk case, in the limit in which membrane fluctuations are unaffected by the wall.

## V Non-equilibrium Membranes: Mode Structure in Bulk

We now discuss the dynamic correlations and mode structure of an impermeable two-component membrane with active force-centres, when placed far away from confining walls. For inactive force centres, the static properties of such a system are given by the free energy of Eq. (11). However, in the presence of non-equilibrium driving forces, thermodynamic restoring forces must be supplemented by additional terms. These are discussed briefly here.

First, we must account for non-equilibrium forces. In the context of the membrane-pump system, these arise due to the action of the pumps. However, since pumping refers to forces internal to the system, the force density exerted on the whole system (membrane $`+`$ pump $`+`$ solvent) must integrate to zero on length scales comparable to the pump size. A simple representation of this requirement idealizes the pump as two force-centres of opposite sign but equal magnitude separated by a distance $`w^{}+w^{}`$ and embedded in the bilayer. Such a “force dipole” representation was introduced in Ref. and will represent an important part of our discussion here.

Second, we must account for the non-equilibrium nature of the noise present in the fluid due to the switching ”on” or ”off” of the pumps. For simplicity, we assume that this noise is delta correlated. Such an assumption corresponds to a coarse-graining in time across the typical pump “dead” time which separates two pumping events. The non-equilibrium active noise can now be absorbed into the correlation function of the equilibrium noise (Eq.(32)). However, note that the “temperature” which appears in this correlator is not the thermodynamic temperature. To smoothly interpolate to the equilibrium case, we must redefine this temperature as an effective temperature which becomes the true thermodynamic temperature in the absence of activity.

Several of the results in this section overlap with those of Manneville et al. (Ref. ) in which the effects of permeation and curvature induced activity on the dynamics of an active membrane in bulk are considered. In our model we assume impermeability from the outset and neglect the effects of curvature induced activity. Such a “minimal” model makes the calculation of the correlation functions of a wall-bounded active membrane more tractable and transparent.

We begin by writing the equations of motion for the various fields. The fluid is incompressible and hence obeys Eq. (33). Since no external forces act on the (fluid+membrane+proteins) system, momentum is conserved  and Eq.(30) holds. The momentum current tensor $`\pi _{ij}`$ is then

$$\pi _{ij}=P\delta _{ij}+\rho v_iv_j\eta (_iv_j+_jv_i)+S_{ij}+A_{ij},$$
(83)
where $`𝐯(𝐫,t)`$ is the three dimensional fluid velocity, $`P(𝐫,t)`$ is the three dimensional pressure field. $`S_{ij}`$ is the contribution to the stress tensor due to membrane conformations and $`A_{ij}`$ is the contribution from the active pumps. Eqns. (31) and (33) can be solved as before for the transverse ($`t`$), longitudinal ($`l`$) and $`z`$ components of the fluid velocity, in terms of the $`t`$,$`l`$ and $`z`$ components of the force $`𝐅(𝐫,t)`$ acting on the fluid due to membrane and pump stresses. The transverse components will be ignored as before; $`𝐟_𝐡(𝐫,t)`$ represents the thermal noise present in the bulk of the fluid.

We can now calculate $`v_z(𝐤_{},0,t)`$ by solving the equations of motion with appropriate boundary conditions. The equation of motion for the height field in the absence of permeation is $`_th(𝐫_{},t)=v_z(𝐫_{},0,t)`$, where $`v_z(𝐫_{},0,t)`$ is the velocity of the surrounding fluid at the mean position of the membrane. The membrane does not feel the forces from the pumps directly because of the absence of permeation but only via the forces the pumps exert on the fluid . The contribution of the bulk equilibrium thermal noise present in the fluid to the $`z`$component of the fluid velocity at $`z=0`$ is $`f_z`$. The statistics of $`f_z`$ have been discussed earlier (see Section III). The density difference field obeys the diffusion equation Eq. (16). As in the equilibrium case, we work in the linearly stable regime.

The membrane fluctuates about $`z=0`$. The force exerted by the membrane and pumps on the fluid is of the form

$$𝐅(𝐤_{},z^{})=\delta (z^{})[\frac{\delta F}{\delta h}\widehat{𝐳}+F_l^m\widehat{𝐤_{}}]+f_{pumpfluid}\widehat{𝐳}.$$
(84)
The force $`f_{pumpfluid}`$ is modelled as a dipolar force density with force-centres located asymmetrically about the $`z=0`$ plane.

$$f_{pumpfluid}=f\psi (𝐫_{},t)[\delta (z^{}w^{})\delta (z^{}+w^{})],$$
(85)
where $`f`$ sets the scale for the active force.

We again solve for the unknown coefficient $`F_l^m`$ using membrane incompressibility. This imposes $`v_l(k_{},0,t)=0`$ in Eq. (34). The hydrodynamic velocity obtained at $`z=0`$ using Eq. (34) enters in the equation of motion for the height field:

$$_th(𝐤_{},t)=\frac{1}{4\eta k_{}}[\frac{\delta F}{\delta h}+f\psi (𝐤_{},t)\mathrm{\Omega }(k_{},w^{},w^{})]+f_z(𝐤_{},0,t),$$
(86)
where

$`\mathrm{\Omega }(𝐤_{},w^{},w^{})`$ $`=`$ $`e^{k_{}w^{}}(1+k_{}w^{})+e^{k_{}w^{}}(1+k_{}w^{})`$ (87)
$`=`$ $`{\displaystyle \frac{k_{}^2}{2}}(w_{}^{}{}_{}{}^{2}w_{}^{}{}_{}{}^{2})+{\displaystyle \frac{k_{}^3}{3}}(w_{}^{}{}_{}{}^{3}w_{}^{}{}_{}{}^{3})+\mathrm{}.`$ (88)
Note that $`\mathrm{\Omega }(k_{},w^{},w^{})`$ vanishes when $`w^{}=w^{}`$, implying that an asymmetry in the position of the force-centres above and below the membrane mid-plane is needed for a non-zero active (non-equilibrium) force. The equations of motion are

$`_t\left(\begin{array}{c}h(𝐤_{},\omega )\\ \psi (𝐤_{},\omega )\end{array}\right)=\left(\begin{array}{cc}\frac{\sigma k_{}^2+\kappa k_{}^4}{4\eta k_{}}& \frac{\kappa \overline{H}k_{}^2f\mathrm{\Omega }(k_{},w^{},w^{})}{4\eta k_{}}\\ \mathrm{\Lambda }\kappa \overline{H}k_{}^4& \mathrm{\Lambda }\chi _0^1k_{}^2\end{array}\right)\left(\begin{array}{c}h(𝐤_{},\omega )\\ \psi (𝐤_{},\omega )\end{array}\right)+\left(\begin{array}{c}f_z(𝐤_{},\omega )\\ i𝐤_{}.𝐟_\psi \end{array}\right).`$ (97)
The correlation function for the height fluctuations of an active membrane (about the flat, stable, steady state phase) is then

$$h^{}(𝐤_{},t)h(𝐤_{},t)=\frac{k_BT}{4\eta k_{}}\frac{(\tau _\psi ^1+\tau _h^1\tau _a^1+\frac{f^2\mathrm{\Omega }^2}{4\eta k_{}\chi _0^1})}{(\tau _\psi ^1+\tau _h^1)(\tau _{h}^{1}{}_{}{}^{e}+\tau _a^1)},$$
(98)
where the quantities $`\tau _\psi ^1,\tau _h^1`$ have been defined earlier. $`\tau _{h}^{1}{}_{}{}^{e}`$ has the same form as $`\tau _h^1`$ with $`\kappa `$ replaced by $`\kappa _e`$. The relaxation time

$$\tau _a^1=\frac{1}{4\eta k_{}}\frac{\kappa \overline{H}}{\chi _{0}^{}{}_{}{}^{1}}f\mathrm{\Omega }(k_{},w^{},w^{})k_{}^2.$$
(99)
We discuss the positivity of the right hand-side of Eq. (98) after discussing the mode structure of the system. From the equal time correlation function we can calculate the quantity $`h^2(L)`$. To lowest order we find

$`h^2(L)=k_BTL^2/\left[\kappa _e{\displaystyle \frac{\kappa \overline{H}𝒫w}{\chi _{0}^{}{}_{}{}^{1}}}\right],`$ (100)
for tensionless membranes and

$`h^2(L)={\displaystyle \frac{k_BT}{\sigma }}\mathrm{ln}(L/a),`$ (101)
for tense membranes. $`𝒫=\frac{f(w_{}^{}{}_{}{}^{2}w_{}^{}{}_{}{}^{2})}{2w}`$, where $`w`$ is the size of the pump, represents the force dipole energy. It is estimated to be a few $`k_BT`$ . These results imply that on large length scales, pumping activity does not change the roughness of equilibrium, tensionless membranes but shifts the temperature at lowest order to

$`T^{eff}=T\left[1+{\displaystyle \frac{\kappa \overline{H}𝒫w}{\kappa _e\chi _{0}^{}{}_{}{}^{1}}}\right].`$ (102)
For tense membranes both roughness and effective temperature are not affected by non-equilibrium effects at large length scales since the divergence of the height correlation function with system size remains logarithmic.

In the stable regime Eq. (100) is positive, as discussed later in this section. The correlation function in Eq. (100) is calculated from the lowest order diffusive terms. Manneville et al.  calculate an identical correlation function but neglect these diffusive terms in favor of higher order terms on the grounds that the diffusion constant is small in the physical situation. For this reason the effective temperature we calculate in Eq. (102) is not the effective temperature that Manneville et al. calculate and compare with experiment.

The height correlation functions we calculate in our linearized approach (Eqns. (100) and (101)) show that the roughness of an active fluid membrane is unchanged in the bulk. The calculation of Ref.  obtains an enhanced roughening. The reason is the different nature of the force densities used in both the calculations. Prost et al. consider a model in which the membrane feels the activity of the pump only as a consequence of permeation. In our calculation, Eq. (86) written as a Langevin equation is

$$_th(𝐤_{},t)+\tau _h^1h(𝐤_{},t)=\frac{(\kappa \overline{H}k_{}^2f\mathrm{\Omega })}{4\eta k_{}}\psi +f_z.$$
(103)
We assume that the pump is always in the active state over the time scales of our interest. The forces on the right-hand side of the above equation arise from three sources: an elastic part (due to the curvature concentration-coupling term in the free energy), a asymmetric dipole part (from the activity of the pumps) and the thermal noise from the fluid. Since the membrane is impermeable it does not feel the active forces or thermal noise directly but only via the fluid. We emphasize that in the present model the membrane senses active forces and thermal noise even when $`\lambda _p=0`$, in contrast to the model of Prost et. al..

The modes are

$`\omega _1,\omega _2`$ $`=`$ $`{\displaystyle \frac{1}{8\eta }}\left(\sigma k_{}+4\eta Dk_{}^2+\kappa k_{}^{}{}_{}{}^{3}\right)`$ (104)
$`\pm `$ $`{\displaystyle \frac{1}{8\eta }}\left(\sqrt{(\sigma k_{}+4\eta Dk_{}^2+\kappa k_{}^{}{}_{}{}^{3})^216\eta (D\kappa _ek_{}^5+D\sigma k_{}^3+f\overline{H}\kappa \mathrm{\Lambda }\mathrm{\Omega }k_{}^3)}\right).`$ (105)
In the absence of curvature-concentration coupling, non-equilibrium effects vanish to lowest order and the modes are those for an equilibrium membrane with hydrodynamic interactions $`\omega _1,\omega _2=(\sigma k_{}+\kappa k_{}^3)/4\eta ,Dk_{}^2`$. When $`\sigma =0`$, curvature-concentration coupling modifies this mode structure to

$`\omega _1`$ $``$ $`{\displaystyle \frac{\kappa _e}{4\eta }}k_{}^3+{\displaystyle \frac{f\kappa \overline{H}(w_{}^{}{}_{}{}^{2}w_{}^{}{}_{}{}^{2})}{8\eta \chi _{0}^{}{}_{}{}^{1}}}k_{}^3,`$ (106)
$`\omega _2`$ $``$ $`Dk_{}^2+{\displaystyle \frac{\kappa \overline{H}(2\kappa \overline{H}f(w_{}^{}{}_{}{}^{2}w_{}^{}{}_{}{}^{2}))k_{}^3}{8\eta \chi _{0}^{}{}_{}{}^{1}}}.`$ (107)
We see that to lowest order, non-equilibrium effects are apparent only at non-zero curvature-concentration coupling. We also observe that the non-equilibrium terms vanish when $`w^{}=w^{}`$.

For stability we require that all the eigenmodes of the system must decay. This means that $`\omega _1`$ and $`\omega _2`$ are positive. To the order above, this imposes $`\frac{\kappa _e}{\kappa }1<\frac{\overline{H}𝒫w}{\chi _0^1}<\frac{\kappa _e}{\kappa }`$. If $`\overline{H}𝒫`$ is such that this condition is satisfied then the system is stable independent of the sign of this parameter. Typical experimental values such as $`\kappa =10k_BT`$, $`\kappa \overline{H}=wk_BT`$,$`\chi _0^1=w^2k_BT`$, $`f=10^{12}N`$, satisfy the above conditions.

## VI Non-equilibrium membranes : Mode Structure in the presence of a wall

We now study the effect of a wall located at $`z=0`$ on the fluctuations of an active membrane located at $`z=d`$. The forces exerted on the fluid by the membrane, pumps and the wall are now given by

$`𝐅(𝐤_{},z^{})`$ $`=`$ $`\delta (z^{}d)[{\displaystyle \frac{\delta F}{\delta h}}\widehat{𝐳}+F_l^m\widehat{𝐤_{}}]+f\psi (𝐫_{},t)[\delta (z^{}dw^{})\delta (z^{}d+w^{})]\widehat{𝐳}`$ (108)
$`+`$ $`\delta (z^{})[F_z^w\widehat{𝐳}+F_l^w\widehat{𝐤_{}}].`$ (109)
We need to determine the unknown coefficients $`F_l^m`$,$`F_z^w`$ and $`F_l^w`$. The following three boundary conditions are used, following Eqns. (34),

$$v_z(𝐤_{},0,t)=v_l(𝐤_{},0,t)=v_l(𝐤_{},d,t)=0.$$
(110)
These boundary conditions impose the following constraints: (i) $`v_z(𝐤_{},z=0)=0`$, i.e. the $`z`$-component of the velocity vanishes at the wall (ii) $`v_l(𝐤_{},z=0)=0`$, i.e. the $`l`$-component of the velocity vanishes at the wall, and (iii) $`v_l(𝐤_{},z=d)=0`$, i.e. the membrane located at $`z=d`$ is incompressible. We neglect the tangential component as before.

The values of the three unknown coefficients are obtained on solving the three simultaneous equations (Eq. (34)) together with the three boundary conditions. These values can then be used to obtain the velocity of the fluid at the location of the membrane. The velocity thus obtained has a part arising from equilibrium elastic forces and a part arising from non-equilibrium active forces. These can be separated out, yielding the equation of motion for the height field as

$`_th(𝐤_{},t)`$ $`=`$ $`v_z(𝐤_{},d,t)`$ (111)
$`=`$ $`𝒢({\displaystyle \frac{\delta F}{\delta h}})+𝒢^{aw}(f\psi )+\stackrel{~}{f_z}(𝐤_{},t),`$ (112)
where $`𝒢`$ is the usual hydrodynamic kernel obtained for the equilibrium fluctuations in the presence of a wall and the new non-equilibrium term

$`𝒢^{aw}(k_{},d,w^{},w^{})`$ $``$ $`\{\begin{array}{cc}\frac{k_{}^2}{12\eta }(3dw_{}^{}{}_{}{}^{2}2w_{}^{}{}_{}{}^{3})+O(k_{}^4)+\mathrm{}\hfill & \text{if }k_{}1/d,\hfill \\ \frac{1}{4\eta k_{}}\mathrm{\Omega }(k_{},w^{},w^{})\hfill & \text{if }k_{}1/d.\hfill \end{array}`$ (115)
is the hydrodynamic kernel for non-equilibrium fluctuations in the presence of the wall(the expanded form of this kernel is given in ; see also Ref.). In the expressions for $`𝒢`$ and $`𝒢^{aw}`$, the lower cut-off on $`k_{}`$ is the inverse of the collision length $`L_c`$, which we will calculate towards the end of this section. The membrane-wall distances we use in our calculations are such that $`d>>w^{},w^{}`$ and hence $`w_{}^{}{}_{}{}^{3}`$ can be neglected in comparison to $`dw_{}^{}{}_{}{}^{2}`$. In the long wavelength limit, $`𝒢^{aw}`$ depends on $`w^{}`$ and not on $`(w^{}w^{})`$ as in the bulk. When the fluctuations are not affected by the wall, $`𝒢^{aw}`$ vanishes when $`w^{}`$ equals $`w^{}`$ since in bulk the directions “up” and “down” are equivalent. The non-equilibrium contribution to the force density therefore vanishes. Thus, in this large membrane-wall distance limit, we recover the previous situation of active membrane fluctuations where the asymmetry was crucial for non-zero active force density. The wall breaks the symmetry in the “up” and “down” directions and hence the hydrodynamic kernel picks out a particular direction and does not depend on $`(w^{}w^{})`$ when membrane fluctuations feel the wall. The statistics for the noise $`\stackrel{~}{f_z}`$ in the presence of the wall follows from Eqns. (79).

The equal time correlation function can be calculated from

$`_t\left(\begin{array}{c}h(𝐤_{},\omega )\\ \psi (𝐤_{},\omega )\end{array}\right)=\left(\begin{array}{cc}𝒢(\sigma k_{}^2+\kappa k_{}^4)& 𝒢\kappa \overline{H}k_{}^2𝒢^{aw}f\\ \mathrm{\Lambda }\kappa \overline{H}k_{}^4& \mathrm{\Lambda }\chi _0^1k_{}^2\end{array}\right)\left(\begin{array}{c}h(𝐤_{},\omega )\\ \psi (𝐤_{},\omega )\end{array}\right)+\left(\begin{array}{c}\stackrel{~}{f_z}(𝐤_{},\omega )\\ i𝐤_{}.𝐟_\psi \end{array}\right).`$ (124)
The correlation function is

$$h^{}(𝐤_{})h(𝐤_{})=k_BT𝒢\left(\frac{(\tau _\psi ^1+\tau _{h}^{1}{}_{}{}^{w}\tau _{a}^{1}{}_{}{}^{w}+\frac{f^2𝒢_{}^{aw}{}_{}{}^{2}}{\chi _0^1𝒢})}{(\tau _\psi ^1+\tau _{h}^{1}{}_{}{}^{w})(\tau _{h}^{1}{}_{}{}^{w,e}+\tau _{a}^{1}{}_{}{}^{w})}\right),$$
(125)
where

$$\tau _{h}^{1}{}_{}{}^{w}=𝒢(\sigma k_{}^2+\kappa k_{}^4),\tau _{a}^{1}{}_{}{}^{w}=𝒢^{aw}\frac{\kappa \overline{H}}{\chi _0^1}fk_{}^2.$$
(126)
$`\tau _{h}^{1}{}_{}{}^{w,e}`$ has the same form as $`\tau _{h}^{1}{}_{}{}^{w}`$ with $`\kappa `$ replaced by $`\kappa _e`$. Inserting physically reasonable values for the parameters which enter these definitions (see below), we verify that the right hand-side of the correlation function is always positive. Thus, this correlation function is calculated in a regime in which the active membrane is in a stable steady-state phase.

At large length scales and in the presence of a confining wall ($`1/L_ck_{}1/d`$),

$$h^2(L)=\frac{k_BT}{\kappa \overline{H}}\left[\frac{ϵ^{}\chi _{0}^{}{}_{}{}^{1}}{ϵf}+\frac{fϵ}{D}\right]ln(L/a)\frac{k_BT}{\mathrm{\Sigma }}ln(L/a),$$
(127)
for tensionless membranes and

$$h^2(L)=k_BT\left[\frac{ϵ^{}+(f^2ϵ^2/D\chi _{0}^{}{}_{}{}^{1})}{ϵ^{}\sigma +(ϵf\kappa \overline{H}/\chi _{0}^{}{}_{}{}^{1})}\right]ln(L/a)\frac{k_BT}{\sigma ^e}ln(L/a),$$
(128)
for tense membranes. We have defined the quantities $`ϵ=dw_{}^{}{}_{}{}^{2}/(4\eta )`$ and $`ϵ^{}=d^3/(12\eta )`$ for notational convenience. Thus, the presence of the wall makes tensionless active membranes less rough than they are in bulk since the height correlation function scales now logarithmically with $`L`$ instead of quadratically. Note that this effect is opposite to that predicted via the scaling arguments of Prost et al. in Ref..

The effective surface tension is then

$`\mathrm{\Sigma }=\kappa \overline{H}/\left[{\displaystyle \frac{ϵ^{}\chi _{0}^{}{}_{}{}^{1}}{ϵf}}+{\displaystyle \frac{fϵ}{D}}\right].`$ (129)
This is an explicitly non-equilibrium effect, since $`\mathrm{\Sigma }=0`$ when $`f=0`$. The induced surface tension also crucially depends on the curvature-concentration coupling $`\overline{H}`$ since it vanishes on setting this coupling to zero. Therefore, both non-equilibrium forces and curvature-concentration coupling are essential for the stiffening of a tensionless, active membrane near a wall.

To calculate the collision length $`L_c`$ we set $`h^2(L_c)d^2`$ in Eqns. (127) and (128). Thus,

$`L_c(d)=\{\begin{array}{cc}aexp\left(\mathrm{\Sigma }d^2/k_BT\right)\hfill & \text{if }\sigma =0,\hfill \\ aexp\left(\sigma ^ed^2/k_BT\right)\hfill & \text{if }\sigma 0.\hfill \end{array}`$ (132)
The eigen-modes have frequencies

$`\omega _1,\omega _2`$ $`=`$ $`{\displaystyle \frac{1}{2}}\left(Dk_{}^2+𝒢\sigma k_{}^2+𝒢\kappa k_{}^{}{}_{}{}^{4}\right)`$ (133)
$`\pm `$ $`{\displaystyle \frac{1}{2}}\left(\sqrt{(Dk_{}^2+𝒢\sigma k_{}^2+𝒢\kappa k_{}^4)^24(f𝒢^{aw}\overline{H}\kappa \mathrm{\Lambda }k_{}^4+𝒢D\kappa _ek_{}^6+𝒢D\sigma k_{}^4)}\right).`$ (134)
In the absence of curvature-concentration coupling, non-equilibrium forces have no effect on the modes and the modes reduce to those of an equilibrium membrane in the presence of a wall, within the linearized theory. For tensionless membranes with non-zero curvature-concentration coupling, non-equilibrium effects become apparent in the mode structure. The modes to lowest order are

$`\omega _1`$ $``$ $`{\displaystyle \frac{f𝒢^{aw}\kappa \overline{H}}{\chi _{0}^{}{}_{}{}^{1}}}k_{}^2+𝒢\kappa _ek_{}^4,`$ (135)
$`\omega _2`$ $``$ $`Dk_{}^2+{\displaystyle \frac{\kappa \overline{H}(\kappa \overline{H}𝒢k_{}^2f𝒢^{aw})}{\chi _{0}^{}{}_{}{}^{1}}}k_{}^2.`$ (136)
When membrane fluctuations are influenced by the wall, $`𝒢=d^3k_{}^2/12\eta `$ and $`𝒢^{aw}=dw_{}^{}{}_{}{}^{2}k_{}^2/4\eta `$, to lowest order. Far away from the wall the modes reduce to those of a non-equilibrium membrane in the bulk.

It may seem that for initially tensionless membranes near a wall, the induced surface tension $`\mathrm{\Sigma }`$ can have either sign depending on the sign of the curvature concentration coupling $`\overline{H}`$. We will show that for stability, it is imperative that $`\overline{H}`$ is positive for active membranes close to walls; a negative $`\overline{H}`$ leads to a negative $`\omega _1`$, and thus an instability. The requirement of positivity of $`\overline{H}`$ in the stable regime ensures that our approximations of considering fluctuations at linear order about an initial flat steady-state phase are self-consistent.

We now present a physical argument for why pumping activity should lead to the stiffening of active, tensionless membranes near repulsive walls. The calculations of Section IV indicate that the eigen-values governing the relaxation of a tense, equilibrium membrane near a wall are $`\sigma d^3k_{}^4/(12\eta )`$ and $`Dk_{}^2`$. For an active, tensionless membrane near a wall these eigenvalues are $`\frac{\kappa \overline{H}fdw_{}^{}{}_{}{}^{2}k_{}^4}{4\eta \chi _0^1}`$ and $`Dk_{}^2`$. The coefficient of the slower mode for the active case ($`O(k_{}^4)`$) can be written as $`\frac{\kappa \overline{H}ϵf}{ϵ^{}\chi _0^1}d^3/(12\eta )`$ and compared to the expression for the slowest relaxation mode for tense, equilibrium membranes. This comparison suggests that an apparent surface tension ($`\frac{\kappa \overline{H}ϵf}{ϵ^{}\chi _0^1}`$) is induced due to purely non-equilibrium effects, even for a membrane with vanishing bare surface tension.

In the linearized theory, the induced surface tension obtained using the above argument and the surface tension $`\mathrm{\Sigma }`$ obtained from the coefficient of the logarithmic term in the calculation of the height roughness coincide. It is thus apparent that the presence of the wall smoothens a tensionless, active membrane by making it relax faster than its tensionless equilibrium counterpart (whose slowest mode is $`O(k_{}^6)`$).

Why does an active membrane relax faster near the wall? Height field fluctuations are predominantly governed by the $`\psi `$ field (the second term in Eq. 111) because this term is $`O(k_{}^2)`$ in contrast to the other terms which are of higher order in wavenumber. Thus, this equation to lowest order in wavenumber is $`_th=𝒢^{aw}f\psi `$. Since the $`\psi `$ field relaxation is fast compared to the height field, we may assume that, over the larger time-scales of interest to us in studying the dynamics of the height field, $`\psi `$ relaxes instantaneously to $`\psi _s=\frac{\kappa \overline{H}}{\chi _{0}^{}{}_{}{}^{1}}k_{}^2h`$. The effective height field equation now becomes

$$_th=\frac{fϵ\kappa \overline{H}}{\chi _{o}^{}{}_{}{}^{1}}k_{}^4h.$$
(137)
Note that the coefficient of the $`k_{}^4`$ term in the above equation can be rewritten to obtain the induced surface tension $`\mathrm{\Sigma }`$, as explained in the previous paragraph.

The above argument invokes and crucially requires the presence of the wall, activity and curvature-concentration coupling. In physical terms, height field fluctuations are accompanied by fast relaxation of the protein density field $`\psi `$. This relaxation is to a steady state value determined by the pump diffusion coefficient and curvature-concentration coupling. The pumping activity is then inhomogeneously distributed along the membrane surface. To see the physical consequences of this inhomogeneous distribution, consider a fluctuation creating a region of positive curvature. If $`\overline{H}`$ is positive, the number of “up” pumps in the region of positive curvature tends to decrease with time and the decrease in the pumping activity causes a smoothening of the membrane. However, if $`\overline{H}`$ is negative, the region of positive curvature attracts more “up” pumps which further bootstraps the height fluctuation, leading to an instability.

The explicit expressions obtained above also permit us to estimate the collision length for tensionless active membranes in the presence of repulsive walls. Using estimates given in for various physical quantities ($`\kappa =10^{20}Joules,\overline{H}=5nm,f=10^{12}N,d=1micron,\chi _{0}^{}{}_{}{}^{1}=10^{21}Joulem^2`$), we find that the induced surface tension $`\mathrm{\Sigma }10^710^8N/m`$. This translates to collision lengths $`L_c`$ of the order of $`10^510^6`$ times molecular scales. Assuming molecular length scales of order a nanometer, this calculation would then predict collision lengths of the order of millimeters, for membranes with physical parameters in the above range.

## VII Conclusions

This paper has presented a calculation of the linearized hydrodynamics of two-component membranes including the effects of curvature-concentration coupling, specifically in the context of a simple model for a biological membrane with protein and lipid constituents. We have studied in some detail the case in which such membranes are placed close to as well as far away from a confining repulsive wall. We have considered the case in which such proteins are “active”, in which case they exert non-equilibrium forces on the solvent, as well as the case of “inactive” pumps, corresponding to thermal equilibrium.

The calculations we report are for a model which shares many similarities with that of Ref.. Manneville et al. consider the more general case of a membrane with a finite permeability whereas we work with an impermeable membrane throughout. In addition, we neglect the possible effects of curvature on activity. However, the relatively simpler structure of our model enables us to establish a connection with equilibrium results in a more transparent fashion. We work in a framework in which boundary conditions are explicitly implemented. This is especially of use when we consider the linearized theory of fluctuations of an active membrane near a wall. The results we report for the linearized hydrodynamics of active and inactive two-component membranes near walls are new, as are our results for the equilibrium case far away from confining walls.

We find, far away from the walls and consistent with the results of Manneville et al. , that the presence of active pumps on the membrane does not change its roughness. In the case of tensionless membranes, fluctuations can be described in terms of an effective temperature which is shifted from the thermodynamic temperature by non-equilibrium effects due to pumping. These terms depend on the quantity $`(w^{}w^{})`$, a measure of the degree of asymmetry in the positions of the force-centres. We reiterate the observation of Manneville et al. in Ref.  that this asymmetry is essential for non-equilibrium effects to be visible in the membrane dynamics.

We find that the activity of the pumps within the stable regime appears to smoothen out height fluctuations in the presence of the wall since height correlations scale only logarithmically with $`L`$ for both tense and tensionless membranes. This smoothening out within the linear theory presumably competes with the possible enhancement of fluctuations, if any, which might arise in a theory which incorporates the effects of non-linearities. We have provided a physical interpretation of this effect in this paper and obtained estimates for the appropriate collision length. We find that this collision length can be significantly larger than typical length scales for biological vesicles, indicating the relevance of the physical effects we study here to a complete description of the hydrodynamics of active pump-membrane systems. It is important to note that this smoothening relies crucially on the existence of curvature-concentration coupling terms, neglected in previous work on the near-wall case. These results suggest that earlier ones based on scaling arguments which predicted an enhancement in the roughness may need to be reevaluated in the light of this work.

We have also calculated the “crossover” function $`𝒢^{aw}`$ and linearized mode structure in the presence of the wall. While the positional asymmetry in the placement of force-centres was crucial for active membrane fluctuations in bulk, we point out that non-equilibrium effects are apparent even for symmetrically placed pumps in the presence of a wall. This unusual, although not totally unexpected, result does not appear to have been noticed before.

We also find that the requirement of stability for active membranes near a wall constrains $`\overline{H}`$, the coefficient coupling concentration with curvature, to be positive. This can have interesting consequences. Consider two active membranes, one with positive $`\overline{H}`$ and the other with negative $`\overline{H}`$, fluctuating in the bulk. Provided $`\overline{H}𝒫`$ for each lies in the window of stability in the bulk, these membranes are stable. Now bring both close to a wall. The membrane with positive $`\overline{H}`$ will remain stable. The fluctuations of the membrane with negative $`\overline{H}`$ however, will turn unstable; its final fate will be decided by the non-linear terms we have omitted from our discussion.

Because our calculation is an explicitly linear one, we cannot address issues such as the relevance of such non-linear terms. The arguments of Prost, Manneville and Bruinsma which predict an anomalous roughening of active membranes near walls rely on identifying the role of such non-linear terms and estimations of their importance via scaling arguments. It would be interesting to construct similar arguments for the model studied here, particularly since our results in the linear case indicate that fluctuations of an active membrane near a repulsive wall can be smoothened in some regimes, rather than roughened relative to the equilibrium case. Further work along these lines is currently in progress.

## Acknowledgments

We are grateful to Sriram Ramaswamy for a critical reading of the manuscript as well as several enlightening discussions on these and related matters. We also thank Y. Hatwalne for urging us to clarify the physical mechanisms underlying our results.

## VIII Appendix

### A  A note on the impermeability of the membrane

In this sub-section we present an argument, following Manneville et. al., to justify the assumption of membrane impermeability. Had we incorporated permeation due to activity in our calculations, active-permeative and active-hydrodynamic terms would appear as $`(\lambda _p\frac{k_{}(w_{}^{}{}_{}{}^{2}w_{}^{}{}_{}{}^{2})}{8\eta })f\psi `$, ($`\lambda _p`$ is the permeation coefficient) . The cross-over length scale from the hydrodynamic to the permeative regime occurs when $`k_{}=\frac{8\eta \lambda _p}{w(w^{}w^{})}`$, i.e. $`l_{}=\frac{\pi w(w^{}w^{})}{4\eta \lambda _p}`$ ($`w`$ is the size of the pump). With $`\lambda _p=10^{12}m^3/N.s`$,$`\eta =10^3kg/m.s`$ and $`w=5.10^9m`$ , this crossover length scale is estimated to be of the order of a centimeter. The length scales of concern to us are in the micron and sub-micron range . Thus, at these length scales, effects due to permeation can be ignored. These arguments can be extended to estimating the role of passive permeative vs. hydrodynamic modes of relaxation; we conclude that at the length scales of interest to us, the assumption of membrane impermeability remains valid.

In the presence of a wall, active permeative terms would dominate if $`\lambda _p>𝒢^{aw}`$. This means that the length scale above which permeation dominates is $`\sqrt{\frac{\pi dw_{}^{}{}_{}{}^{2}}{2\eta \lambda _p}}`$. This length is approximately $`40`$ microns when the membrane-wall distance $`d=1`$ micron. Hence, in the regimes of interest to us, permeation can be ignored and relaxation is purely hydrodynamic. Hence, we assume $`\lambda _p=0`$ to derive our results.

### B A modified model for active membranes

We present here results for a model in which proteins are located asymmetrically across the bilayer with their centers of mass displaced above or below the bilayer midpoint (see Fig. 3). The head and tail of the pumps are then located at distances $`w_l`$ and $`w_s`$ respectively from the midpoint of the bilayer. Note that in this model, the “up” and “down” pumps are related to each other by reflection along the $`z=0`$ line unlike the model of Fig. 2. This model can be used to describe situations in which proteins are attached to planar bilayer membranes which are symmetric on both sides, unlike vesicles in which the inner leaf and outer leaf need not be symmetrically preferred by the protein.

When pumps are active on such a membrane in bulk, the force density due to the pumps acting on the fluid has the form

$`f_{pumpfluid}=f[\delta (zw_l)n^{}\delta (z+w_s)n^{}\delta (z+w_l)n^{}+\delta (zw_s)n^{}],`$ (138)
where $`n^{}`$ and $`n^{}`$ are the local number of “up” and “down” pumps respectively. On repeating the calculations done in Section V, we find that the $`z`$-component of the hydrodynamic velocity has the same form as shown in Eq. (5.7) with $`\mathrm{\Omega }(k_{},w^{},w^{})`$ replaced by $`\mathrm{\Omega }^{}(k_{},w_l,w_s)=e^{k_{}w_s}(1+k_{}w_s)+e^{k_{}w_l}(1+k_{}w_l)`$, which vanishes when $`w_l=w_s`$ (i.e. for symmetrically placed proteins). Correlation functions and relaxation times can be read off from the corresponding expressions in Section V by identifying $`w^{}`$ and $`w^{}`$ with $`w_s`$ and $`w_l`$ respectively. For example,

$$h^{}(𝐤_{},t)h(𝐤_{},t)=\frac{k_BT}{4\eta k_{}}\left(\frac{(\tau _\psi ^1+\tau _h^1\tau _a^1+\frac{f^2\mathrm{\Omega }^2}{4\eta k_{}\chi _0^1})}{(\tau _\psi ^1+\tau _h^1)(\tau _{h}^{1}{}_{}{}^{e}+\tau _a^1)}\right),$$
(139)
where the quantities $`\tau _\psi ^1,\tau _h^1`$ have been defined earlier. $`\tau _{h}^{1}{}_{}{}^{e}`$ has the same form as $`\tau _h^1`$ with $`\kappa `$ replaced by $`\kappa _e`$. The relaxation time

$$\tau _a^1=\frac{1}{4\eta k_{}}\frac{\kappa \overline{H}}{\chi _{0}^{}{}_{}{}^{1}}f\mathrm{\Omega }(k_{},w_l,w_s)k_{}^2.$$
(140)
We note here that this identification is purely notational since $`(w^{},w^{})`$ and $`(w_s,w_l)`$ physically represent different lengths in the two models.

When the calculations are repeated for such a model in the presence of a repulsive wall, we find that the hydrodynamic velocity acquires an additional contribution from the $`\varphi `$ (total protein density) field. In this linearized calculation we assume that the compressibility associated with the $`\varphi `$ field is very large and ignore its dynamics. However, we do not expect this contribution to alter the long wavelength fluctuations of the active membrane near the wall significantly.

Figures