# Physical Parameters for Biconcave Shape Vesicles

## Equation and Requirements

For an axisymmetric surface $`\mathrm{\Sigma }`$, we take the rotational axis to be $`z`$-axis and the plane of reflection $`xy`$-plane, then a biconcave shape is obtained by revolving and reflecting a curve, given by a function $`z(r)0`$ defined for $`r`$ in some interval $`[0,r_{\mathrm{}}]`$, where $`r_{\mathrm{}}`$ is the radius of the “equator”. For a biconcave axisymmetric surface, we expect $`z(r)`$ and its derivative $`w=z^{}`$ to have the following graphs.

Usually, the Helfrich shape equation of axisymmetrc vesicles is written in term of the angle $`\psi `$ between the surface tangent and the plane perpendicular to rotational axis . For our discussion, it is convenient to rewrite it in terms of the derivative of the graph $`w(r)=z^{}(r)=\mathrm{tan}\psi `$. Then the Helfrich shape equation of axisymmetrc vesicles becomes an equation for $`w`$, which is

$`{\displaystyle \frac{2rw^{\prime \prime }}{(1+w^2)^{5/2}}}`$ $`={\displaystyle \frac{5rww_{}^{}{}_{}{}^{2}}{(1+w^2)^{7/2}}}`$ (1)   
$`{\displaystyle \frac{2w^{}}{(1+w^2)^{5/2}}}+{\displaystyle \frac{2w+w^3}{r(1+w^2)^{3/2}}}`$   
$`+{\displaystyle \frac{2c_0w^2}{1+w^2}}+{\displaystyle \frac{(c_0^2+\stackrel{~}{\lambda })rw}{(1+w^2)^{1/2}}}{\displaystyle \frac{\stackrel{~}{p}r^2}{2}}.`$   
The biconcave shape and symmetries are then translated into suitable initial and boundary conditions, such as, $`w(0)=0`$; $`w^{}(0)=w_0^{}>0`$; $`w(r)\mathrm{}`$ as $`rr_{\mathrm{}}`$; an integral condition that $`\mathrm{}{\displaystyle _0^r_{\mathrm{}}}w\mathrm{d}r<0`$; and $`w`$ is required to have a unique local maximum and no other critical points.

The lower order terms can be given by polynomials $`Q(t)`$ or its quadratic part $`R(t)=Q(t)t^3`$. After multiplying with $`rw^{}`$, the equation (1) can be written as follow,

$$\begin{array}{cc}\hfill \left[\frac{r^2w_{}^{}{}_{}{}^{2}}{(1+w^2)^{5/2}}\right]^{}& =\left[\frac{w^2}{(1+w^2)^{1/2}}\right]^{}\hfill \\ & +r^3w^{}R(\kappa (r));\hfill \end{array}$$
(1a)   
$$\begin{array}{cc}\hfill \left[\frac{r^2w_{}^{}{}_{}{}^{2}}{(1+w^2)^{5/2}}\right]^{}& =\left[\frac{2}{\sqrt{1+w^2}}\right]^{}\hfill \\ & +r^3w^{}Q(\kappa (r)).\hfill \end{array}$$
(1b)   
where $`\kappa (r)={\displaystyle \frac{w}{r\sqrt{1+w^2}}}`$ is the principal curvature of $`\mathrm{\Sigma }`$ in the meridinal direction. Indeed,

$$\kappa (r)w_0^{},\text{as }r0\text{,}$$

so the initial choice $`w_0^{}`$ is the central meridinal curvature or $`w_{0}^{}{}_{}{}^{2}`$ the Gaussian curvature. Note that if $`c_0=\stackrel{~}{\lambda }=\stackrel{~}{p}=0`$, equation (1a) can be integrated directly to a solution $`z=z(r)`$ which is a circular arc with radius $`1/w_0^{}`$. This is exactly the case when the meridinal curvature is a constant.

## Necessary Conditions

The lower order terms in (1) capture the changes of the meridinal curvature. Since $`Q(0)=R(0)=\stackrel{~}{p}/2<0`$, if $`w_0^{}`$ is less than the smallest positive root of $`Q(t)`$, then both $`Q(w_0^{})<0`$ and $`R(w_0^{})<0`$. Our analysis shows that these negativity conditions are essential for the meridinal curvature $`\kappa (r)`$ to change sign. Furthermore, if the physical parameters constitute a polynomial $`Q`$ with only positive roots, a solution $`w(r)`$ with $`w_0^{}`$ small enough always satisfies all of our requirements. The threshold value of $`w_0^{}`$ can be calculated numerically if exact values of the parameters are given.

The meridinal curvature $`\kappa (r)`$ of the vesicle $`\mathrm{\Sigma }`$ is the most basic quantity in our analysis. From equation (1b), if $`c_0>0`$ and the initial curvature at the center $`w_0^{}`$ is large, in the sense that $`R(w_0^{})>0`$, then $`\kappa (r)`$ increases and a biconcave shape cannot be formed. So a necessary condition for formation of axisymmetric solution of biconcave shape in the case that $`c_0>0`$ is

$$R(w_0^{})=2c_0w_{0}^{}{}_{}{}^{2}+(c_0^2+\stackrel{~}{\lambda })w_0^{}\frac{\stackrel{~}{p}}{2}<0.$$
(2)   
The following pictures show the behaviors of $`\kappa (r)`$ for two different values of $`w_0^{}`$ with $`c_0=1`$, $`\stackrel{~}{\lambda }=0.25`$, and $`\stackrel{~}{p}=1`$. For these parameter values, $`R(0.277124)=0`$.

$`R\left(w_0^{}\right)>0`$ 

$`R\left(w_0^{}\right)<0`$ 

The one on the left is taken with $`w_0^{}=0.278`$ so the necessary condition is not satisfied. In this case, $`\kappa `$ blows-up at finite radial distance and the solution only leads to a convex surface. The picture on the right is taken with $`w_0^{}=0.276`$. In this case, the necessary condition is satisfied, so $`\kappa `$ decreases and changes sign. But it only decays gradually and the integral area condition is not satisfied. As a result, the surface and its reflection intersects in $`^3`$.

There is a more subtle boundary condition imposed by the reflection symmetry. One needs to verify that $`z(r)`$ and $`z(r)`$ together form a stationary solution with respect to any variation near the plane of reflection. For the situation that we discuss, it turns out that this is true. Moreover, we obtain a relation between the radius $`r_{\mathrm{}}`$ of the “equator” and the Gaussian curvature $`K(r_{\mathrm{}})`$ of the vesicle at any point on the “equator”,

$$K(r_{\mathrm{}})^2=\frac{1}{r_{\mathrm{}}}Q\left(\frac{1}{r_{\mathrm{}}}\right).$$
(3)   
This relation is easily verified for the case that $`c_0=\stackrel{~}{\lambda }=\stackrel{~}{p}=0`$, in which, $`Q(t)=t^3`$ and $`K(r_{\mathrm{}})^2={\displaystyle \frac{1}{r_{\mathrm{}}^4}}`$. This is compatible with the fact that the solution surface is a round sphere.

## Sufficient condition

To have a solution meeting all the requirements, we need to impose a stronger condition that $`w_0^{}`$ to be small enough, at least smaller than the positive roots of $`Q`$. This smallness assumption on $`w_0^{}`$ implies that not only $`R(w_0^{})<0`$ but also $`Q(w_0^{})<0`$. Both $`\kappa (r)`$ and $`w(r)`$ start initially positive and become negative when $`r`$ reaches a certain value $`r_0`$ (depending on the value of $`w_0^{}`$, actually, comparable to $`w_{0}^{}{}_{}{}^{1/2}`$ when $`w_0^{}`$ is small). In fact, we discover that, for any point of inflection $`r_c`$ of the graph of $`z(r)`$ with $`\kappa (r_c)>0`$,

$$\frac{16}{3\stackrel{~}{p}}\underset{w_0^{}0}{lim}\frac{r_c^2}{w_0^{}}\underset{w_0^{}0}{lim}\frac{r_0^2}{w_0^{}}\frac{16}{\stackrel{~}{p}}.$$

As a consequence of these inequalities, the graph of $`z(r)`$ has a unique point of inflection. After $`r`$ reaches $`r_0`$, with the assumption that all roots of $`Q(t)`$ are positive, the lower order terms of equation (1b) are dominated and the function $`w(r)`$ blows down monotonically to $`\mathrm{}`$ in a finite radial distance. By further estimating the area according to $`w_0^{}`$, the integral area condition can be verified whenever $`w_0^{}`$ is sufficiently small. All these behaviors of the functions $`z(r)`$ and $`w(r)`$ with respect to $`Q`$ and the choice of $`w_0^{}`$ are illustrated in our summary on p. Discussions. The mathematical details will be presented in a forthcoming paper.

## Discussions

Finally, we would like to remark on other solutions which do not correspond to biconcave surfaces.

The one on the left is a solution for $`c_0=1`$, $`\stackrel{~}{\lambda }=3`$, $`\stackrel{~}{p}=2`$ with $`w_0^{}=2`$. Note that $`w_0^{}`$ is greater than the positive roots of $`R(t)`$ and $`Q(t)`$. Moreover, $`Q(t)`$ has negative roots and thus this is beyond the situation discussed above. In this case, after $`w=z^{}`$ becomes negative, it may not decrease monotonically to negative infinity, which is why a second “petal” may form.

The other type is a rotational solution which is not symmetric under reflection in $`xy`$-plane . The picture is produced with $`c_0=1`$, $`\stackrel{~}{\lambda }=1`$ and $`\stackrel{~}{p}=1`$. The “upper” curve satisfies our equation (1) but not the area condition. The “lower” curve and the reflected “upper” curve are both solutions locally, so there is a bifurcation of solution at $`r_{\mathrm{}}`$. The surface only closes up by an asymmetric one. It should be remarked that, from our numerical study, not all solutions in the upper quadrant may be closed up by an asymmetric counterpart. For example, if the upper part satisfies the area condition, the only possible lower part is the reflection symmetric one. This suggests a possible uniqueness of biconcave solution. However, at the moment, there is not yet a mathematical proof.

| Graph of $`Q`$ and positions of $`w_0^{}`$ ($`c_0=1`$, $`\stackrel{~}{\lambda }=0.25`$, $`\stackrel{~}{p}=1`$) |  |  |
| --- | --- | --- |
| Graphs of solution $`z`$ | | |
| (a) $`w_0^{}=0.4`$ | (b) $`w_0^{}=0.25`$ | (c) $`w_0^{}=0.18`$ |
|  |  |  |

| Graph of $`Q`$ and positions of $`w_0^{}`$ ($`c_0=1`$, $`\stackrel{~}{\lambda }=0.3`$, $`\stackrel{~}{p}=1`$) |  |  |
| --- | --- | --- |
| Graphs of solution $`z`$ | | |
| (a) $`w_0^{}=0.4`$ | (b) $`w_0^{}=0.27`$ | (c) $`w_0^{}=0.15`$ |
|  |  |  |

| Graph of $`Q`$ and positions of $`w_0^{}`$ ($`c_0=1`$, $`\stackrel{~}{\lambda }=0.25`$, $`\stackrel{~}{p}=1`$) |  |  |
| --- | --- | --- |
| Graphs of solution $`z`$ | | |
| (a) $`w_0^{}=2`$ | (b) $`w_0^{}=0.5`$ | (c) $`w_0^{}=0.275`$ |
|  |  |  |