How are Saturating Removal (SR, Uri Alon et al.) and Minimal Aging model (Fedichev & Gruber) related?

I recently read the following paper by Uri Alon and his group:

https://doi.org/10.64898/2025.12.22.695887

It's a great paper! They study their own aging model (Saturating Removal) as to how different parameters affect healthspan and lifespan. In addition, they also do a similar analysis for the Minimal Aging model by Peter Fedichev and Jan Gruber. The nice thing about the paper is that they are very thorough in documenting how they compute things and give explicit values for the model parameters after fitting them to a dataset or when presenting figures. Very nice!

Further, the paper has a set of supplemental material, which covers some of the analytical properties of both models.

Reading this paper deepened a notion I already had before. Namely that both models seem to be very similar. In this little article I want to investigate how similar they actually are.

1. Mathematical definition of both models

Let's first remind ourselves of the definition of both aging models.

Note: Parameters that appear in both models with the same letter, will have a suffix SRor FGto indicate their origin.

1.1. SR model

The SR model is described by the following differential equation,

\[ \frac{dX}{dt} = ηt - β_{\text{SR}} \frac{X}{X + κ} + \sqrt{2ε}ξ(t), \]

where "η,βSR,κ,ϵare model parameters, while ξrepresents Gaussian white noise. η has units of time⁻², βSRof time⁻¹, εof time⁻¹, κis normalized to 1, and Xand Xcare in units of κ." (from the supp. mat.). Note that "κis normalized to 1" means that it is a dimensionless constant, not that its value is exactly 1 (in practice it typically seems to be κ=0.5).

In the model, ηis the damage accumulation parameter. It is linear in time, which means the damage accumulation rate increases linearly with time. This is a modeling of the aging process, in which the large number of failing subsystems result in ever more damage appearing at faster rates.

βSRis the maximum removal capacity of damage (hence the negative sign of the term). At small Xvalues, the term is effectively βSRX/κ, which means the removal rate is proportional to the existing damage (i.e. fast healing rates etc.). At large Xvalues, XX+κ1, which means a fixed damage removal rate of βSR. κis the amount of damage Xat which the relative removal rate is half that of the total removal capacity and thus represents an important point at which the nonlinear saturation property starts to become important. Note that this is early in life / at very low total damage values. A typical death threshold is Xc17.5, while κis often κ=0.5.

The last term with parameter εis the Gaussian noise the system is under at all times.

1.2. F&G model

In contrast, the Fedichev & Gruber model is defined by

\begin{align} \frac{dz_0}{dt} &= β_{\text{FG}}Z(t) - (ε_0 - β'Z(t)) z_0 + gz²_0 + J_0 + \sqrt{2D_0} ξ \\ \frac{dZ}{dt} &= γ \end{align}

The fast variable z0represents a reversible “resilience” or leading physiological state of the organism. The slow variable \(Z(t) = γ t + Z_0\) is the irreversible accumulation of entropic damage, where γis the damage accumulation rate (DAR).

The parameters are interpreted as follows: βFG couples damage production to the slow damage level, ε0 is the baseline linear restoring force that pulls z0back toward equilibrium, β'quantifies how accumulating damage Z gradually erodes this resilience, gis the strength of the quadratic destabilizing nonlinearity, J0is an optional constant external stress, and D0sets the amplitude of Gaussian noise.

Note that the second equation has a simple solution. It is solved by

\[ Z(t) = γt + Z_0 \]

where Z0is some initial damage.

If we insert this into the first differential equation, we get the following form,

\begin{align} \label{eq:fg_expanded} \frac{dz_0}{dt} &= β_{\text{FG}}(γt + Z_0) - (ε_0 - β'(γt + Z_0)) z_0 + gz²_0 + J_0 + \sqrt{2D_0} ξ \\ &= β_{\text{FG}}γt + β_{\text{FG}}Z_0 - (ε_0 - β'γt - β'Z_0) z_0 + gz²_0 + J_0 + \sqrt{2D_0} ξ \\ &= (β_{\text{FG}} + β'z_0) γt + β_{\text{FG}}Z_0 - (ε_0 - β'Z_0) z_0 + gz²_0 + J_0 + \sqrt{2D_0} ξ. \\ \end{align}

In the second equation I've grouped everything by powers of z0 and in the last equation by terms based on γt.

Importantly, death in the F&G model is not defined by a hard threshold on z0(in contrast to SR’s Xc). Instead, death occurs via loss of dynamical stability: as Z(t)increases, the effective restoring force ε0β'Z(t)weakens until a saddle-node bifurcation is reached (the stable fixed point disappears). Mortality then arises from stochastic escape over the flattening potential barrier. This stability-based death criterion is a direct mathematical consequence of the quadratic truncation, as we will see in sec. 10.

2. Generic Langevin potential differential equation

Both models represent a differential equation for a particle in a potential well, undergoing stochastic random noise and are a Langevin equation.

In a generic sense, we can write

\begin{equation} \label{eq:generic_langevin} \frac{d x}{d t} = A t + B - C x + G x² + \sqrt{2 D} ξ(t), \end{equation}

where we have a direct time dependence with coefficient A, a constant coefficient B, a negative linear term in x, a quadratic term in x2and the stochastic random noise term with coefficient 2D, where I have written the time dependence of the random process explicitly. The latter is what makes this a stochastic differential equation (and the omission of the explicit time dependence in both papers is imo annoying).

When comparing the two models above with this equation, we can see that they sort of follow this equation, but not quite:

  • the SR model does not seem to have the terms Cx+Gx2,
  • the F&G model has a term β'z0γtin addition, which would correspond to something like Ttxin our notation here (with a new parameter T).

In the next few sections we will show that these differences are to a large extent cosmetic.

3. SR model: linear and quadratic terms

The form presented in eq. \eqref{eq:generic_langevin} does not seem to show up in the SR model. The equivalent of the xvariable in the SR model is the Xvariable. Cx+Gx2contains all terms that are proportional to xin some sense. Thus, the corresponding term in SR, is

\[ -β_{\text{SR}}\frac{X}{X + κ}. \]

Now, if you are a physicist like me or someone with more applied math experience, you will probably realize that these two are more similar than they appear at a first glance. Consider this:

  • at very small Xvalues where Xκ, the fraction behaves like a linear (!) function
  • at slightly larger values the value of κbecomes more important. This slows the growth of the linear curve. This can be modeled by a negative quadratic term.

To make this more concrete, we can do the Taylor expansion 1 of this fraction around X=0,

\[ \frac{X}{X + κ} = \frac{X}{κ} - \frac{X²}{κ²} + \frac{X³}{κ³} - \frac{X⁴}{κ⁴} + \mathcal{O}(X⁵). \]

If we ignore all terms of X3or higher, this is exactly the form of the generic Langevin equation.

Applying this to the SR model as a whole yields the following,

\begin{equation} \label{eq:sr_model_taylor} \frac{dX}{dt} = ηt - β_{\text{SR}} \frac{X}{κ} + β_{\text{SR}}\frac{X²}{κ²} + \sqrt{2ε}ξ. \end{equation}

This is eq. \eqref{eq:generic_langevin} with

\begin{align} A &= η & B &= 0 & C &= \frac{β_{\text{SR}}}{κ} \\ G &= \frac{β_{\text{SR}}}{κ²} & D &= ε & & \end{align}

Later in sec. 10 we will discuss the validity of this Taylor expansion in more detail. In particular how it relates to the F&G model, the death criteria and the death threshold Xcin SR.

4. F&G model: an additional tz0term

The F&G model in expanded form eq. \eqref{eq:fg_expanded} is,

\[ \frac{d z_0}{dt} = β_{\text{FG}}γt + β_{\text{FG}}Z_0 + J_0 - (ε_0 - β'Z_0) z_0 + gz²_0 + β'γt z_0 + \sqrt{2D_0} ξ \]

This lets us identify the equivalent coefficients compared to the generic Langevin equation \eqref{eq:generic_langevin} as,

\begin{align} A &= β_{\text{FG}}γ & B &= β_{\text{FG}}Z_0 + J_0 & C &= ε_0 - β'Z_0 \\ G &= g & D &= D_0 & & \end{align}

which only leaves us with one mismatched term. The β'γtz0term is a direct coupling between the time and position variables.

This leaves us with the initial conclusion of restricting us to the case of β'=0. With that restriction the F&G model reduces to,

\begin{equation} \label{eq:fg_beta_zero} \frac{d z_0}{dt} = β_{\text{FG}}γt + β_{\text{FG}}Z_0 + J_0 - ε_0 z_0 + gz²_0 + \sqrt{2D_0} ξ \end{equation}

5. Preliminary comparison of SR and F&G model

Now compare the Taylor expanded version of the SR model, eq. \eqref{eq:sr_model_taylor} with the F&G variant without explicit time coupling, eq. \eqref{eq:fg_beta_zero}. We can then conclude the two models are equivalent if and only if,

\begin{align} η &= β_{\text{FG}}γ & Z_0 &= J_0 = 0 & β_{\text{SR}} &= ε_0 \\ \frac{β_{\text{SR}}}{κ²} &= g & ε &= D_{0} & & \end{align}

In other words, if we accept zero starting entropy Z0and no constant stress J0in the F&G model and restrict the parameter space of ε0to ε0=gFGκ2SR(and thus reducing a degree of freedom as ε0and gare coupled via the κparameter), the two models are identical for small X.

6. Allow constant offset in SR model

In order to relax the restriction that Z0=J0=0to obtain the same model as SR, we can make a minor extension to the SR model.

Namely, by performing a time shift \(t ↦ t + τ\), we gain an additional parameter, which can capture the constants. In general terms, the differential equation is,

\[ \frac{dX}{dt} = f(t, X(t)), \]

where fis the placeholder for the exact definition, which is solved by X(t). If we define a new solution,

\[ Y(t) \coloneq X(t + τ), \]

we arrive at,

\[ \frac{dY}{dt} = \frac{d}{dt} X(t + τ) = f(t + τ, X(t + τ)) = f(t + τ, Y(t)). \]

That means, the time shifted equation with \(t ↦ t + τ\) is solved by Y(t). Writing out this equation explicitly in our case,

\begin{equation} \label{eq:sr_time_shifted} \frac{dX}{dt} = η(t + τ) - β \frac{X}{κ} + β\frac{X²}{κ²} + \sqrt{2ε}ξ. \end{equation}

At this point the term ητ=Bcorresponds to the constant offset. This time shifted version of the SR model can thus be mapped to the F&G model via either,

  • ητ=Z0,J0=0,
  • ητ=Z0+J0
  • or ητ=J0,Z0=0

depending on preference or purpose.

7. Mapping F&G to SR

At this point we can map the F&G model to the SR model in an almost general fashion (with the only limitation still being β'=0).

If we map the coefficients of eq. \eqref{eq:generic_langevin} from the SR model with constant offset and the F&G model and solve them for the SR parameters, we arrive at the following mapping:

\begin{align} \label{eq:mapping_fg_sr} η &= β_{\text{FG}}γ & β_{\text{SR}} &= \frac{ε_0²}{g} \\ τ &= \frac{1}{γ}\left( Z_0 + \frac{J_0}{β_{\text{FG}}}\right) & κ &= \frac{ε_0}{g} \\ ε &= D_0 & & \end{align}

This allows to map any specific instantiation of the F&G model to an equivalent SR model.

Note that the τparameter, is independent of a. This makes a lot of sense, as the new time parameter τis just an offset of "starting damage" / "initial entropy". It does not depend on how damage accumulates over time or generally the system evolves.

8. Finally: introduce time-position coupling in SR model

We have now seen that with some minor modifications to the SR model, the two are almost identical. The one mismatch is the existence of the β'parameter in the F&G model, which couples its z0variable to its tvariable. We may wish to recover the lost degree of freedom from setting β'=0in the F&G model.

By taking βto be β(t)in the SR model in addition to the previous modifications, we can then also recover the coupling of time and position. In the simplest form we may take,

\[ β(t) = β_0 + β_1 t \]

We now plug this into our time shifted variant of the SR model, eq. \eqref{eq:sr_time_shifted}.

\begin{align} \frac{dX}{dt} &= η(t + τ) - β(t) \frac{X}{κ} + β(t)\frac{X²}{κ²} + \sqrt{2ε}ξ \\ &= η(t + τ) - β_0 \frac{X}{κ} + β_0 \frac{X²}{κ²} - β_1 t \frac{X}{κ} + β_1 t \frac{X²}{κ²} + \sqrt{2ε}ξ \end{align}

If we map β0to the normal βwe had before we now have two additional terms, β1tXκ+β1tX2κ2. The former is the equivalent of the β'z0γtterm in the F&G model. We are justified in ignoring the latter term, as the product tX2is of a higher order than the terms we decided to include after our Taylor expansion, which we cut off at X3. This means our final model is,

\begin{equation} \frac{dX}{dt} = η(t + τ) - β_0 \frac{X}{κ} + β_0 \frac{X²}{κ²} - β_1 t \frac{X}{κ} + \sqrt{2ε}ξ \end{equation}

9. Fully generic Langevin equation including time coupling

If we include a time position coupling into the general Langevin equation \eqref{eq:generic_langevin} from above, we arrive at this equation with added parameter T:

\begin{equation} \label{eq:generic_langevin_with_time} \frac{d x}{d t} = A t + T t x + B - C x + G x² + \sqrt{2 D} ξ(t). \end{equation}

We can use this to map both the F&G model and SR model to these now six parameters.

It leads to the following expressions, which while appearing slightly more complicated are a logical extension of the mapping in sec. Mapping F&G to SR, eq. \eqref{eq:mapping_fg_sr}:

\begin{align} \label{eq:full_mapping_fg_sr} η &= β_{\text{FG}}γ & β_{\text{SR}} &= \frac{(ε_0 - β'Z_0)²}{g} \\ τ &= \frac{1}{γ}\left( Z_0 + \frac{J_0}{β_{\text{FG}}}\right) & κ &= \frac{(ε_0 - β' Z_0)}{g} \\ ε &= D_0 & β_1 &= -\frac{ε_0 - β' Z_0}{g} β' γ \end{align}

Essentially, we need to replace ε0by ε0β'Z0and we have an additional equation for the new parameter β1.

This is our final parameter mapping. This is an almost fully general mapping between the SR and F&G models, including all parameters of the F&G model. This mapping maps all eight parameters of the F&G model to the now six parameters of the SR model.

10. Validity range of the Taylor approximation and implications for the two models

Having discussed the mapping of our modified SR model to the F&G model, let's now come back to one of the biggest modifications / assumptions we made: the Taylor expansion of the XX+κ term in the SR model. As mentioned previously, this Taylor expansion is valid for small values of X, because we expanded around \(X = 0\). Small here is relative to the magnitude of κ, which tends to be at κ=0.5. However, the SR model defines death as the damage crossing the critical damage threshold Xc17.5. Those kind of damage values are well outside the validity range of the Taylor expansion.

Fig. 1 compares the relative damage removal rate of the actual term XX+κwith the first three orders of the Taylor expansion, linear Xκ, quadratic Xκ+X2κ2and additionally cubic Xκ+X2κ2X3κ3. We can see that for values up to around κ0.2the three expansions are roughly compatible. However, beyond that they start to diverge rapidly with the linear and cubic terms exploding, indicating the damage removal keeping up with the damage, while the quadratic term actually goes back down to zero (and technically towards ).

sr_taylor_expansion_validity.svg
Figure 1: SR saturating removal vs. Taylor expansion over the full biologically relevant range. Exact removal rate βXX+κ(red line), linear approximation (green), quadratic truncation that recovers the F&G form (cyan), cubic for reference (purple). Parameters: κ=0.5(typical), β=1(normalization). Vertical dashed line: typical death threshold Xc17.5. The inserted plot is the same zoomed to the small-X regime, up to X=1.

This limited validity range directly explains several apparent differences between the two models and highlights their complementary strengths.

Crucially, the SR model yields sensible results that reproduce biological systems up to large damage values. In contrast, a model like F&G or our modified SR model does not behave sensibly at large damage values. SR thus has more robust global behavior.

While this may appear to point to an enormous problem in our mapping between the models, the (in my opinion) correct interpretation is a lot more interesting. The fact that the Taylor expanded version is not sensible for large values of Ximplies the F&G model must use a different definition of death out of necessity. So while the two models can look very similar at the level of the differential equation, their actual application to an 'agent simulation' is inherently different.

11. Summary & conclusion

Effectively, the SR and F&G models are similar models, with some different choices driven by their different definitions. Viewing the F&G model as a local approximation of the SR model explains why the F&G model cannot have a global damage threshold as the definition for death. At the same time, the F&G model in practice contains more independent parameters and includes constant offsets.

The functional definition of SR's differential equation contains F&G as a local, small-Xapproximation. The additional parameters that F&G exposes can only be modeled after mild modification (time shift, time dependent β(t)).

Footnotes:

1

A Taylor expansion of a function is a representation of said function around a specific point via an infinite series of polynomials. In physics it is very often used to reduce a complex expression to simpler, lower order terms that are accurate for small values, by cutting off the expansion at a fixed order.

Author: basti

Created: 2026-02-23 Mon 11:20

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