text:
# Application of Statisitcal Physics to Politics

## 1 Setting the limits

Modern theory of critical phenomena is based on the fundamental concepts of universality and irrelevant variables . These two concepts mean that different physical systems, like for instance a magnet and a liquid, behave the same way when passing from one macroscopic state to another macroscopic state. Well known examples are the magnet becoming a para-magnet, the liquid, a gas, and even may be the creation of the universe from nothing with the big bang. Most of the microscopic properties of the physical compounds involved turn out to be irrelevant for describing the macroscopic change which in turn appears to be universal. While the number of physical systems undergoing phase transitions is infinite, all associated phase transitions can be described in terms of only a small finite number of universality classes. Only a few parameters, like space dimensionality, determine which universality class the system belongs to. The abstract and general nature of the statistical physics framework makes it tempting to extend such notions to non-physical systems, and in particular to social systems, for which, in many cases, there exists an interplay between microscopic properties and macroscopic realities.

Nevertheless the two fields of physical sciences and social sciences are rather different. However, the process of going in parallel from one atom and one human to respectively several atoms in bulk and a social group has much in common. More precisely, it is the hypothesis behind the present approach. It is worth stressing that we are not claiming our models will explain all aspects of human behavior. Like any modeling effort, it is appropriate only to some classes of phenomena in social science and not to others.

However such an approach should be carefully controlled. To just map a physical theory built for a physical reality, onto a social reality, could be at best a nice metaphore, but without predictability, and at worst a misleading and wrong social theory. Physics has been successfull in describing macroscopic behavior using properties of the constituant microscopic elements. The task here, is to borrow from physics those techniques and concepts used to tackle the complexity of aggregations. The challenge is then to build a collective theory of social behavior along similar lines, but within the specific constraints of the psycho-social reality. The constant danger is for the theorist to stay in physics, using a social terminology and a physical formalism. The contribution from physics should thus be restricted to qualitative guidelines for the mathematical modeling of complex social realities. Such a limitation does not make the program less ambitious.

## 2 Real space: from physics to politics

In this paper we present an application of statistical physics to political sciences . We apply the concept and technics of real space renormalization group  to study the majority rule voting process within hierarchical structures. In particular we focus on the conditions for a given political party to get for sure, full power at the hierarchy top level.

We find that majority rule voting produces critical threshold to total power. Having an initial support above the critical threshold guarantees full power at the top. The value of the critical threshold to power is a function of the voting structure, namely the size of voting groups and the number of voting hierarchical levels. Using these results a new explanation is given to the sudden and abrupt fall of eastern former communist parties.

Here we apply the real space renormalization group scheme of collective phenomena in physics to a radically different reality, namely a social one We emphasis on other technical aspects than the ones usually used in physics. While there it is an abstract and formal tool to study phase transitions, here we associate a political reality to each step of the renormalization group transformation. On this basis it is worth to stress we apply statistical physics to political sciences not as a qualitative metaphor but indeed as a guide to build a quantitative model to study the effect of majority rule voting on hte democratic representation of groups within a hierarchical organization.

Like in physics we start from local cells constituted by a small number of degrees of freedom, here individuals. Similarly to Ising spins, to keep the analysis simple, people can choose only between two political tendencies A and B. Associated proportions of A and B support in the system, a political group, a firm, a network, a society, are supposed to be known. We denote them by $`p_0`$ for the overall A-support and $`1p_0`$ for the B-support. We are assuming each member does have an opinion.

Once formed, each cell elects a representative, either an A or a B using a majority rule. These elected people (the equivalent of the super spin rescaled to an Ising one in real space renormalization group) constitute the first hierarchical level of the organization called level-1. New cells are then formed at level-1 from these elected people. They in turn elect new representatives to build level-2. This process is repeated again and again.

In physics the rescaled degree of freedom is fictius while here it is a real person. We are not using a theoretical scheme to embody some complex features but instead we are building up a real organization where each voting step is real. Moreover at odd with physics the number of iterations in the renormalization process are finite and the focus is on the stable fixed points.

The following of the paper is as follows. In next section we present and study the case of 3-person cells. A critical threshold to power is singled out. It equals $`50\%`$. A minority is found to self-eliminate within a few voting levels. A making sense bias in the voting rule is then introduced in Section 3. A one vote bonus is added to the tendency already in power in cases of A-B equality. For 4-person cells, this bias shift the critical threshold to power from $`50\%`$ to $`77\%`$. Size effects are analyzed in Section 4. Analytic formulas are then derived. In particular, given an initial support $`p_0`$ for the A, the number of voting levels necessary to their self-elimination is obtained. Section 5 puts the results in a more practical perspective. Last section contains a discussion about both former apparent eternity of communist leaderships as well as their sudden collapse. Some.perspectives are outlined.

## 3 The simplest fair case

We start from a population distributed among two tendencies A and B with respectively $`p_0`$ and $`1p_0`$ proportions within the the system. It could be either a political group, a firm, or a whole society. At this stage each member does have an opinion. From now on we will use the political language.

Cells are constituted by randomly aggregating group of 3 persons. It could be by home localization or working place. Each cell then elects a representative using majority rule. To have an A elected requires either 3 A or 2 A in the cell. Otherwise it is a b who is elected. these elected persons constitute the first level of the hierarchy denoted level-1. The same process of cell forming can be repeated within level-1 from the elected persons. Making the cell to vote produces an additional level, namely level-2. We can go on the same way from a level-n to a level-(n+1). The probability to have an A elected at level (n+1) is then,

$$p_{n+1}P_3(p_n)=p_n^3+3p_n^2(1p_n),$$
(1)
where $`p_n`$ is the proportion of elected A persons at level-n.

We call $`P_3(p_n)`$ the voting function. It has 3 fixed points $`p_d=0`$, $`p_{c,3}=\frac{1}{2}`$ and $`p_t=1`$. First one corresponds to the disappearance of the A. Last one $`p_t`$ represents the totalitarian situation where only A are present. Both are stable. At contrast $`p_c`$ is unstable. It determines the threshold to full power. Starting from $`p_0<\frac{1}{2}`$ repeating voting leads towards 0 while the flow is in direction of 1 for $`p_0>\frac{1}{2}`$.

Therefore majority rule voting produces the self-elimination of any proportion of the A-tendency as long as $`p_0<\frac{1}{2}`$. However the democratic self-elimination to be completed requires a sufficient number of voting levels.

At this stage the instrumental question is to determine the number of levels required to ensure full leadership to the initial larger tendency. The analysis will turn relevant to reality only if this level numbers is small. Most organizations has only a few level, and always less than 10.

For instance starting from $`p_0=0.43`$ we get successively $`p_1=0.40`$, $`p_2=0.35`$, $`p_3=0.28`$, $`p_4=0.20`$, $`p_5=0.10`$, $`p_6=0.03`$ down to $`p_7=0.00`$. Therefore 7 levels are sufficient to self-eliminate $`43\%`$ of the population.

Tough the aggregating voting process eliminate a tendency it stays democratic since it is the leading tendency (more than $`50\%`$ ) which after all gets the total leadership of the organization. It is worth to notice the symmetry with respect to A nd B tendencies.

## 4 The simplest killing case

In real world things are not as fair as above and often it turns out very hard, if not impossible, to change an organization leadership. We will now illustrate this situation.

Considering yet the simplest case we constitute groups of 4 persons instead of 3. The salient new feature is the 2A-2B configuration for which there exists no clear majority. In most social situations it is well admitted that to change a policy required a clear cut majority. In case of no decision, things will stay as they are. It is a bias in favor of the already existing. Often this bias is achieved giving for instance, one additional vote to the committee president.

Along this line, the voting function becomes non symmetrical. Assuming the B were in power, for an A to be elected at level $`n+1`$ we have,

$$p_{n+1}P_4(p_n)=p_n^4+4p_n^3(1p_n),$$
(2)
where $`p_n`$ is the proportion of elected A persons at level-n. At contrast for a B it is,

$$1P_4(p_n)=p_n^4+4p_n^3(1p_n)+2p_n^2(1p_n)^2,$$
(3)
where last term embodies the bias in favor of B. From Eqs (2) and (3) the stable fixed points are still 0 and 1. However the unstable one is drastically shifted to,

$$p_{c,4}=\frac{1+\sqrt{13}}{6},$$
(4)
which makes the threshold to power to the A about $`77\%`$. Moreover the process of self-elimination is accelerated. For instance from $`p_0=0.69`$ we have the series $`p_1=0.63`$, $`p_2=0.53`$, $`p_3=0.36`$, $`p_4=0.14`$, $`p_5=0.01`$, and $`p_6=0.00`$. It shows that using an a priori reasonable bias in favor of the B turns a majority rule democratic voting to a totalitarian outcome. Indeed to get to power the A must pass over $`77\%`$ of support which almost out of a possibility. Above series shows how $`63\%`$ of a population disappears from the leadership within only 5 voting levels.

## 5 Larger voting groups

Most real organizations work with voting cells larger than 3 or 4. To account for this size variable we generalize above scheme to cells with r voting persons. We then have to determine the voting function $`p_{n+1}=P_r(p_n)`$. Using a majority rule it becomes,

$$P_r(p_n)=\underset{l=r}{\overset{l=m}{}}\frac{r!}{l!(rl)!}p_n^l(1+p_n)^{rl},$$
(5)
where $`m=\frac{r+1}{2}`$ for odd r and $`m=\frac{r+1}{2}`$ for even r which thus accounts for the B-bias.

The two stable fixed points $`p_d=0`$ and $`p_t=1`$ are preserved with enlarging the group size. However while the unstable one stays $`p_{c,r}=\frac{1}{2}`$ for odd sizes, it varies with r for even sizes. It starts at $`p_{c,4}=\frac{1+\sqrt{13}}{6}`$ with the limit $`p_{c,r}\frac{1}{2}`$ when $`r\mathrm{}`$.

We can then calculate analytically the critical number of levels $`n_c`$ at which $`p_{n_c}=ϵ`$ with $`ϵ`$ being a very small number. It determines the level of confidence of the prediction to have no A elected.. One way to evaluate $`n_c`$ is to expand the voting function $`p_n=P_r(n1)`$ around the unstable fixed point $`p_{c,r}`$,

$$p_np_{c,r}+(p_{n1}p_{c,r})\lambda _r,$$
(6)
where $`\lambda _r\frac{dP_r(p_n)}{dp_n}|_{p_{c,r}}`$ with $`P_r(p_c)=p_{c,r}`$. Rewriting last equation as,

$$p_np_{c,r}(p_{n1}p_{c,r})\lambda _r,$$
(7)
we can then iterate the process to get,

$$p_np_{c,r}(p_0p_{c,r})\lambda _r^n,$$
(8)
from which we get the critical number of levels $`n_c`$ at which $`p_n=ϵ`$. Taking the $`\mathrm{ln}`$ on both side of Eq. (8) gives,

$$n\frac{\mathrm{ln}(p_cp_0)}{\mathrm{ln}\lambda _r}+n_0,$$
(9)
where $`n_0\frac{\mathrm{ln}(p_{c,r}ϵ)}{\mathrm{ln}\lambda _r}`$ is only valid not too far from $`p_{c,r}`$. However it turns out to be a rather good estimate even down to the sable fixed point 0 by making it equal to 1 while taking the integer part of Eq .(9). For a more accurate calculation of $`n_0`$ see .

## 6 The magic formula

Most organizations don’t change their structure at every election or decision event. They are set once and then don’t change any longer. The number of hierarchical levels is thus fixed and constant. Therefore to make our analysis useful we have to invert the question on“how many levels are needed to eliminate a tendency” onto “given n levels what is the necessary overall support to get full power”.

It is worth to keep in mind that situations for respectively A and B tendencies are not always symetric. Here we stress the dynamics of voting with respect to the A. To implement theis operative question, we invert Eq.(7) to,

$$p_0=p_{c,r}+(p_np_{c,r})\lambda _r^n.$$
(10)
indeed two critical thresholds now appears. First one, the disappearence threshold $`p_{d,r}^n`$ which gives the value of support under which the A disappears for sure at the top level leadership. It is given by Eq. (10) putting $`p_n=0`$,

$$p_{d,r}^n=p_{c,r}(1\lambda _r^n).$$
(11)
In parallel putting $`p_n=1`$ give the second threshold $`p_{f,r}^n`$ above which the A get full and total power. Using Eq.(11), we get,

$$p_{f,r}^n=p_{d,r}^n+\lambda _r^n.$$
(12)
which shows the appearence of a new region for $`p_{d,r}^n<p_0<p_{f,r}^n`$. In that region the A neither disappear totally nor get full power ($`p_n`$ is neither $`0`$ nor $`1`$). It is therefore a coexistence region where some democraty is prevailing since results of the election process are only probabilistic. No tendency is sure of winning making alternating leadership a reality. However as seen from Eq.(12), this democratic region shrinks as a power law $`\lambda _r^n`$ of the number n of hierarchical levels. Having a small number of levels thus puts higher the threshold to a total reversal of power but simultaneously lowers the threshold for non existence.

Again, above formulas are approximates since we have neglected corrections in the vicinity of the stable fixed points. However they give the right qualitative behavior. Actually $`p_{d,r}^n`$ fits to $`n+1`$ and $`p_{f,r}^n`$ to $`n+2`$. For more accurate formulas see .

To get a practical feeling of what Eqs. (11) and (12) means, let us illustrate them for the case $`r=4`$ where we have $`\lambda =1.64`$ and $`p_{c,4}=\frac{1+\sqrt{13}}{6}`$. Considering 3, 4, 5, 6 and 7 level organizations, $`p_{d,r}^n`$ is equals to respectively $`0.59`$, $`0.66`$, $`0.70`$, $`0.73`$ and $`0.74`$. In parallel $`p_{f,r}^n`$ equals $`0.82`$, $`0.80`$, $`0.79`$, $`0.78`$ and $`0.78`$. These series emphasizes drastically the totalitarian character of the voting process.

## 7 Some prospective

Up to now we have treated very simple cases to single out main trends produced by democratic voting aggregating over several levels. In particular we have singled out the existence of critical thresholds to full power. Moreover these tresholds are not necesseraly symetric for both tendencies in competition. In the biased 4-cell case it is around $`0.77\%`$ for the A.

Such asymetries are indeed always present in most realistic situations in which more than two groups are competing. Let us consider for instance the case of three competing groups A, B and C. Assuming a 3-cell case, now the ABC configuration is unsolved using majority rule as it was for the precedent two tendencies AABB 4-cell configuration. For the AB case we made the bias in favor of the group already in power, like giving an additional vote to the comitee president.

For multi-group competitions typically the bias results from parties agreement. Usually the two largest parties, say A and B are hostile while the smallest one C would compromise with either one of them. Then the ABC configuration gives a C elected. In such a case, we need 2A or 2B to elect respectively an A or a B. Otherwise a C is elected. Therefore the elective function for A and B are the same as for the AB 3-cell model. It means that the critical threshold to full power to A and B is $`50\%`$. In otherwords for initial A and B supports less than $`50\%`$ the C get full power. The required number of levels is obtained from above formulas.

It is possible to generalize to as many groups as wanted. The analysis becomes more heavy but the mean features of voting flows towards fixed point are preserved.

## 8 Conclusion

To conclude we comment on some possible new explanation to the recent generalized auto-collapse of eastern communist parties. Up to this historical and drastic event, communist parties seemed eternal with the same leadership ever. Once they collapsed most explanations were related to both an opportunistic change within the various organizations together with the end of the soviet army threat.

May be the explanation is different and related to our hierachical model. Indeed communist organizations are based on the structutral concept of democratic centralism which nothing else than a tree-like hierachy with a rather high critical threshold to power. Suppose it was of order of $`80\%`$ like in our 4-cell case. We could than consider that the internal opposition to the orthodox leadership did grew a lot and massively over several decades to evantually reach and pass the critical threshold with the associated sudden outrise of the internal opposition. Therefore the at once collapse of eatern communist pparties would have been the result of a very long and solid phenomena. Such an explanation does not oppose to additional constraints but emphazie on the internal mechanism within these organizations.

At this stage it is of importance to stress that modeling social and political phenomena is not stating some absolute truth but insted to single out some basic trend within very complexe situations.


format: text

text:
# Stochastic Resonance in Neuron Models: Endogenous Stimulation Revisited

## I Introduction

The improvement of signal transmission and detection through noise has been studied keenly over the past two decades under the paradigm of stochastic resonance; for recent reviews see Refs. . This holds true for the neurosciences in particular, which have for a long time been puzzled by the seemingly irregular activity of the nervous system. A long series of experiments has now firmly established that sensory neurons of various modalities benefit from ambient noise .

Theoretical efforts to explain stochastic resonance in neuronal systems had to abstract rigorously from the biological complexity of real neurons to facilitate mathematical treatment . The leaky integrate-and-fire model neuron (LIF)  is likely the most widely studied of these abstract neuron models, especially in investigations of the neuronal code . The main advantages of this model are its simplicity and lack of memory: Each time the neuron has been excited sufficiently to fire a spike (output pulse), it is reset to a predefined state, erasing all memory of past input. One may thus analyze the intervals between spikes separately, and build the complete spike train fired by the neuron by concatenation of intervals.

Time-dependent stimulation complicates this procedure, though, as a different stimulus is now presented during each interspike interval. This precludes a straightforward analysis of global properties of the spike train, such as its power spectral density. Since stochastic resonance is typically defined in terms of the signal-to-noise ratio in response to periodic stimulation, either of the following is required: (i) further simplification of the model, or (ii) the development of better techniques.

The latter is certainly preferable and has been achieved in the meantime . Stochastic resonance with respect to both noise amplitude and signal frequency has been found, and may be relevant to signal processing in neuronal networks . Results obtained for the leaky integrate-and-fire neuron are in qualitative agreement with those obtained from more realistic, non-linear models , indicating that the abstraction to the LIF model was acceptable.

Albeit stochastic resonance in the leaky integrate-and-fire model may thus be considered a solved problem, we would like to return here to some earlier studies , which, at least in parts, followed the simplifying approach (i). These studies assumed that the same stimulus was presented during each interspike interval, i.e., that the originally periodic stimulus was reset to a fixed phase $`\varphi _0`$ after each spike. All intervals are thus equivalent, and global properties of the spike train may be computed from the interspike-interval distribution for a single interval using the methods of renewal theory . Lánský coined the terms *endogenous* and *exogenous* stimulation for stimulation with and without reset, respectively .

This stimulus-reset assumption is patently unbiological for virtually all neurons, since it would require the neuron to fully control the input it receives. The reset assumption was therefore justified as an approximation to the full, exogenous dynamics of a periodically driven noisy neuron along the following lines : If the neuron is driven by a periodic subthreshold stimulus (a stimulus too weak to evoke spikes in the absence of noise), then the neuron will fire all spikes at approximately the same stimulus phase $`\varphi ^{}`$ after transients have died out. This phase corresponds roughly to the phase at which the membrane potential is maximal in the absence of noise. Such firing patterns are found in sensory neurons, e.g., in the auditory nerve  or cold-receptor neurons . Thus, roughly the same stimulus, starting from phase $`\varphi (\tau =0)\varphi ^{}`$, is presented during each interval. Resetting the stimulus phase to a fixed phase $`\varphi (\tau =0)=\varphi _0`$ will therefore introduce only minor errors.

This reasoning suffers from an essential shortcoming, namely the choice of a *fixed* reset phase $`\varphi _0`$ independent of both stimulus and noise. This reset phase is a free parameter of the model, which has no counterpart in biological neurons. We show here that stochastic resonance in terms of the signal-to-noise ratio as studied previously in the LIF model with stimulus reset  occurs because the model neuron adapts best to the free parameter “reset phase” for a particular noise amplitude. The signal-to-noise ratio *as defined in those studies* diverges monotonically for vanishing noise if the reset phase is adapted to stimulus and noise in a plausible way.

The leaky integrate-and-fire model and the methods applied in analysis are briefly reviewed in Sec. II; see Ref.  for details. Results are presented in Sec. III and summarized in Sec. IV.

## II Model and Methods

### A Leaky integrate-and-fire neuron

The leaky integrate-and-fire neuron model sketches the neuron as a capacitor with leak current, which is charged by an input current $`I(t)`$ until the potential $`v(t)`$ across the capacitor (membrane potential) reaches a threshold $`\mathrm{\Theta }`$. At that instant, an output spike is recorded and the potential reset to $`v_r<\mathrm{\Theta }`$. The assumption that the stimulus is reset after each spike as well implies that the membrane potential evolves according to

$$\dot{v}(\tau )=v(\tau )+\mu +q\mathrm{cos}(\mathrm{\Omega }\tau +\varphi _0)+\sigma \xi (\tau ),$$
(1)
in between any two spikes ; $`\tau `$ is intra-interval time, i.e., $`\tau `$ runs from zero in every interval. $`\mu `$ is the DC component of the stimulus, $`q`$ its AC amplitude, $`\mathrm{\Omega }`$ the nominal frequency, and $`\varphi _0`$ the fixed but arbitrary reset phase. All quantities are measured in their natural units, i.e., the membrane time constant $`\tau _m`$ and threshold $`\mathrm{\Theta }`$. $`v_r=0`$ is assumed throughout. The noise term in Eq. (1) subsumes both biochemical and network noise , and is taken to be Gaussian white noise \[$`\xi (t)\xi (t^{})=\delta (tt^{})`$\]. A different realization of the noise is used for each interval. Sub-threshold stimuli are characterized by $`sup_t\mathrm{}v(t)=\mu +q/\sqrt{1+\mathrm{\Omega }^2}<1`$ for $`\sigma =0`$. We restrict ourselves here to these stimuli, because they appear to be more relevant for the encoding of periodic signals in sensory systems than superthreshold stimuli .

The sequence $`\tau _1,\tau _2,\mathrm{},\tau _k,\mathrm{}`$ of intervals corresponds to an output spike train $`f(t)=_k\delta (tt_k)`$ with spike times $`t_k=_{jk}\tau _j`$. This spike train is evoked by an *effective stimulus* consisting of piecewise sinusoids of length $`\tau _k`$, as shown in Fig. 1. In contrast, we call the pure sinusoid $`\mathrm{cos}(\mathrm{\Omega }t+\varphi _0)`$ the *nominal stimulus*. Figure 1 indicates that the effective stimulus approximates the nominal stimulus for a reset phase of $`\varphi _00`$, while it is an irregular sequence of piecewise sinusoids for other choices of the reset phase. We therefore focus here on $`\varphi _0=0`$ in accordance with earlier work .

Because of the stimulus reset and the whiteness of the noise, all interspike interval lengths $`\tau _k`$ are statistically independent, identically distributed random variables with density $`\rho (\tau )`$. The latter can be computed numerically or approximated in closed form . The sequence of intervals thus forms a renewal process, which is fully characterized by the ISI density $`\rho (\tau )`$ . Periodic sub-threshold stimulation evokes multimodal ISI densities as shown in Fig. 2 for noise not too strong. The location of the first peak depends on the reset phase $`\varphi _0`$, while subsequent peaks follow at intervals of the nominal stimulus period $`T=2\pi /\mathrm{\Omega }`$. Comparable ISI distributions are found in sensory neurons .

### B Signal-to-noise ratio

The performance of a signal processor is commonly measured in terms of the signal-to-noise ratio in studies on stochastic resonance. Since the spike train elicited from the neuron is a renewal process by virtue of the stimulus reset, its power spectral density (PSD) is given by

$$S(\omega )=\frac{1}{\pi \tau }\left(1+2\mathrm{}\frac{\stackrel{~}{\rho }(\omega )}{1\stackrel{~}{\rho }(\omega )}\right),\omega >0,$$
(2)
where

$$\stackrel{~}{\rho }(\omega )=_0^{\mathrm{}}\rho (\tau )e^{i\omega \tau }𝑑\tau $$
(3)
is the Fourier transform of the ISI density and $`\tau `$ the mean interspike interval length; note that $`\rho (\tau )=0`$ for $`\tau <0`$ by definition.

The input to the neuron is not purely sinusoidal due to the stimulus reset, and the maximum of the PSD will thus be shifted away from the stimulus frequency $`\mathrm{\Omega }`$, see Fig. 3(a). We thus define the signal as the maximum of the PSD in a window around $`\mathrm{\Omega }`$

$$\widehat{S}=S(\widehat{\mathrm{\Omega }})=\mathrm{max}\{S(\omega )|0.9\mathrm{\Omega }<\omega <1.1\mathrm{\Omega }\}$$
(4)
and refer to the location $`\widehat{\mathrm{\Omega }}`$ of the maximum as *peak frequency*. The signal $`\widehat{S}`$ is undefined if $`S(\omega )`$ has no absolute maximum within the window as, e.g., in Fig. 3(b). The white power spectrum of a Poissonian spike train of equal intensity, $`S_P=(\pi \tau )^1`$, is used as reference noise level , whence the signal-to-noise ratio is

$$R_{\text{SN}}=\widehat{S}/S_p=\pi \tau \widehat{S}.$$
(5)
Note that power spectral density and signal-to-noise ratio as defined above are calculated for infinitely long spike trains. Any strictly periodic component of the spike train will thus give rise to singularities in the PSD. In the reset model, coherence is broken by the stimulus reset, resulting in a continuous spectrum . Due to the different definitions, signal-to-noise ratios obtained from the LIF model with and without stimulus reset cannot be compared quantitatively .

The spectrum defined by Eq. (2) may have very narrow peaks for low noise, whence numerical evaluation of Eqs. (2)–(5) may require very high frequency resolution.

### C Preferred firing phase

If the stimulus is not reset after each spike, the probability $`\chi ^{(k+1)}(\psi )`$ for the $`k+1`$-st spike to occur at stimulus phase $`\psi `$ can be expressed in terms of the corresponding distribution for the $`k`$-th spike as

$$\chi ^{(k+1)}(\psi )=_\pi ^\pi 𝒯(\psi |\varphi )\chi ^{(k)}(\varphi )𝑑\varphi $$
(6)
where $`𝒯(\psi |\varphi )`$ is a stochastic kernel . For $`k\mathrm{}`$, the firing phase distribution will approach a stationary distribution $`\chi ^{(s)}`$ and we choose the preferred firing phase as the phase at which the neuron most likely fires,

$$\varphi ^{}=\mathrm{arg}\underset{\psi }{\mathrm{max}}\chi ^{(s)}(\psi ).$$
(7)
Using this preferred phase as reset phase will yield a viable approximation to stimulation without reset only if the firing phase distribution is sharply concentrated around $`\varphi ^{}`$, and has only a single maximum. To ensure this, we require that the vector strength  of the distribution fulfills

$$r=\left|e^{i\varphi }\right|=\left|_\pi ^\pi e^{i\varphi }\chi ^{(s)}(\varphi )𝑑\varphi \right|0.9.$$
(8)
This condition is generally met by the sub-threshold stimuli combined with weak noise studied here. Multimodal firing phase distributions are observed only for slow stimuli in the presence of intermediate to large noise, and for superthreshold stimuli. We are therefore not concerned with complications that may arise from mode-locking as observed in the latter cases .

## III Results

### A Fixed Reset Phase

For fixed reset phase, $`\varphi _0=0`$, the model neuron shows typical stochastic resonance behavior, i.e., a maximum of the SNR at an intermediate albeit small noise amplitude $`\sigma _{\text{max}}`$ as shown in Fig. 4(a) . The mechanism inducing stochastic resonance is indicated in Fig. 4(b): the maximal SNR is reached when the peak frequency $`\widehat{\mathrm{\Omega }}`$ and the reset frequency $`\mathrm{\Omega }_{\text{res}}=2\pi /T_{\text{res}}`$ coincide, where $`T_{\text{res}}`$ is the mode of the ISI density, i.e., the most probable interval between two stimulus resets. Coincidence of reset and peak frequencies thus indicates synchronization between the stimulus reset and the correlations dominating the power spectrum of the output spike train, yielding optimal encoding of the nominal stimulus.

This effect may intuitively be explained as follows. Assume that the peaks of the ISI distribution are located around $`\tau =nT+ϵ`$, $`ϵ>0`$, for a given choice of reset phase and small noise. The effective stimulus will then be a close to the nominal stimulus, cf. Fig. 1(a). The neuron will fire at shorter intervals as the noise is increased, and for a particular noise amplitude will the peaks of the ISI distribution be centered about multiples of the stimulus period $`T`$. The effective stimulus will then be reset in intervals of $`nT`$. The neuron is thus on average driven by a periodic stimulus with period $`\mathrm{\Omega }_{\text{res}}=\mathrm{\Omega }`$, evoking as periodic a spike train as possible, i.e., one maximizing the signal-to-noise ratio. This explanation ignores all jitter in spike timing, which reduces the SNR. The SNR maximum is thus not found for that input noise amplitude which yields $`\mathrm{\Omega }_{\text{res}}=\mathrm{\Omega }`$ ($`\sigma 0.018`$), but for somewhat weaker input noise, corresponding to smaller output jitter, cf. Fig. 4(b).

### B Noise-Adapted Reset Phase

A LIF neuron driven by a sinusoidal stimulus which is *not* reset after each spike will approach a stationary firing pattern . The preferred firing phase in the stationary state is given by Eq. (7) and depends on the noise amplitude as shown in Fig. 5(c): the neuron fires at later phases for weaker noise. Note that interspike intervals will be multiples of the stimulus period $`T`$ in this regime, since the neuron fires all spikes at the same phase $`\varphi ^{}`$ (up to jitter).

This observation suggests how to construct a proper approximation to the full LIF dynamics using the reset model: for each stimulus and each noise amplitude, determine the preferred phase from Eq. (7) and use this phase as reset phase,

$$\varphi _0=\varphi _0(\sigma ;\mu ,q,\mathrm{\Omega })=\varphi ^{}(\sigma ;\mu ,q,\mathrm{\Omega }).$$
(9)
The stimulus is then reset in intervals of multiples of the stimulus period, so that the effective stimulus will differ from the nominal stimulus only through jitter. This jitter vanishes as input noise vanishes, whence perfect periodic stimulation will be attained for $`\sigma 0`$. Consequently, the signal-to-noise ratio will diverge for vanishing noise as shown in Fig. 5(a). The reset frequency is identical to the nominal frequency by construction, $`\mathrm{\Omega }_{\text{res}}=\mathrm{\Omega }`$, cf. Fig. 5(b). The peak frequency, on the other hand, converges to the stimulus frequency as noise vanishes, $`\widehat{\mathrm{\Omega }}\mathrm{\Omega }_{\text{res}}=\mathrm{\Omega }`$, as the effective stimulus becomes identical to the nominal one.

We shall now make this argument rigorous. The ISI distribution of a LIF model neuron is well approximated by

$$\rho (\tau )h(\tau )\mathrm{exp}\left[_0^\tau h(s)𝑑s\right]$$
(10)
$$h(\tau )=\mathrm{exp}\left[\left(\frac{1v_0(\tau )}{\sigma }\right)^2\right]$$
(11)
where $`v_0(\tau )`$ is the membrane potential in the absence of noise, i.e., the solution of Eq. (1) for $`\sigma =0`$. We consider the limit of small noise ($`\sigma 1`$) and slow stimulation ($`T1`$). In that case, we can neglect transient terms in the membrane potential to obtain

$$v_0(\tau )=\mu +\widehat{q}\mathrm{cos}(\mathrm{\Omega }\tau +\mathrm{\Omega }\zeta )+𝒪(e^\tau ),$$
(12)
where $`\widehat{q}=1/\sqrt{1+\mathrm{\Omega }^2}`$, and $`\zeta =(\varphi _0\mathrm{arctan}\mathrm{\Omega })/\mathrm{\Omega }`$. The hazard $`h(\tau )`$ will thus be a sequence of narrow peaks around the maxima of $`v_0(\tau )`$, i.e. around $`\tau _n=nT\zeta `$. ISI lengths are multiples of the stimulus period for adapted reset phase by construction, whence $`\zeta =0`$ for $`\varphi _0=\varphi ^{}(\sigma )`$.

As peaks are narrow, we may take the exponential term in Eq. (10) to be constant across each peak; it merely reduces the peak amplitude by a factor $`\gamma `$ for each subsequent peak. The ISI distribution can thus be approximated as a superposition of dampened copies of the hazard function centered about $`\tau _n=nT`$, i.e.,

$$\rho (\tau )\underset{n=1}{\overset{\mathrm{}}{}}\gamma ^{n1}\mathrm{exp}\left[\left(\frac{1v_0(\tau \tau _n)}{\sigma }\right)^2\right]$$
(13)
with $`\gamma =\mathrm{exp}\left(_{T/2}^{3T/2}h(s)𝑑s\right)`$. Expanding the exponent separately for each summand and retaining only terms of lowest order in $`\tau \tau _n`$ yields,

$$\rho (\tau )c\underset{n=1}{\overset{\mathrm{}}{}}\gamma ^{n1}\mathrm{exp}\left(\frac{\mathrm{\Omega }^2(\tau nT)^2}{\eta \sigma ^2}\right),$$
(14)
where $`\eta ^1=\widehat{q}(1\mu \widehat{q})`$, and $`c`$ is a normalization factor. The interspike-interval distribution has thus been reduced to a sum of Gaussians. This approximation holds well for small noise, as shown in Fig. 6.

The Fourier transform of the ISI distribution is thus

$$\stackrel{~}{\rho }(\omega )=\frac{(1\gamma )\mathrm{exp}\left[\frac{\eta \sigma ^2}{4}\left(\frac{\omega }{\mathrm{\Omega }}\right)^2+2\pi i\frac{\omega }{\mathrm{\Omega }}\right]}{1\gamma \mathrm{exp}\left[2\pi i\frac{\omega }{\mathrm{\Omega }}\right]}.$$
(15)
In the limit of small noise and for adapted reset phase, the effective stimulus becomes equal to the nominal stimulus, and the spectral power will be maximal at the nominal stimulus frequency in this limit, i.e., $`\widehat{\mathrm{\Omega }}\mathrm{\Omega }`$ for $`\sigma 0`$, cf. Fig. 5(b). The signal-to-noise ratio is therefore

$$R_{\text{SN}}=\frac{S(\mathrm{\Omega })}{S_P}\mathrm{coth}\frac{\eta \sigma ^2}{8}(\sigma 1)$$
(16)
in good agreement with numerical results for the exact model (solid line in Fig. 5(a)). We find in particular that the signal-to-noise ratio diverges for vanishing noise in the Gaussian approximation.

## IV Conclusions

The aim of this paper was to clarify whether the response of the leaky integrate-and-fire neuron to periodic subthreshold stimulation can be approximated as a renewal process. In particular, we wanted to know whether stochastic resonance at weak noise as reported in earlier work  is a genuine property of the LIF neuron, or rather an artefact of the stimulus reset, which was introduced to reduce the full neuronal dynamics to a renewal process.

We argued that renewal (endogenous) dynamics are a good approximation to the full (exogenous) dynamics only if the phase $`\varphi _0`$, to which the stimulus is reset after each spike, is adapted to the stimulus parameters, especially the noise amplitude, in such a way that the neuron fires most likely at phase $`\varphi _0`$. We showed that stochastic resonance does not occur in this case. Stochastic resonance is only found if the reset phase is held fixed as the amplitude of the input noise is varied.

There is no biologically plausible way in which a neuron could reset a stimulus impinging on it to a fixed phase. Lánský suggested that neurons driven by internally generated membrane potential oscillations could reset their oscillator to a fixed phase, hence the term endogenous stimulation . Cold receptor cells are driven by internal subthreshold oscillations, but do not reset their internal oscillator upon firing . It is thus highly unlikely to find neurons which reset the oscillator that drives them to a fixed phase independent of stimulus properties, i.e., follow genuine endogenous dynamics. This in turn means that any effect arising solely from the reset to a fixed stimulus phase will not be found in real neurons. The stochastic resonance effect reported in Ref.  (see also ) is thus a model artefact.

We stress that this conclusion applies only to the particular type of stochastic resonance discussed here. The leaky integrate-and-fire neuron benefits from stochastic resonance when encoding periodic signals *without* reset (exogenous stimulation), as shown in . This resonance occurs at noise amplitudes which are about one order of magnitude larger than the resonance found in the model with reset to a fixed phase. The crucial difference is that studies on exogenous stimulation consider either explicitly or implicitly the power spectral density of a spike train of finite duration, while an infinite spike train is assumed here, cf. Sec. II B. Trains of very low intensity, but precisely phase-locking to the stimulus, as found for very weak noise, yield a small finite-time signal-to-noise ratio, while their infinite-time SNR may be large.

Other noise-induced resonance phenomena found in the LIF with stimulus reset, e.g., in the interspike-interval distribution or the mean ISI length , are not directly affected by our finding either. Indeed, Shimokawa et al. found comparable resonance effects in terms of the ISI distribution for stimulation with and without reset . These resonances occur at much larger noise amplitudes than studied here; see also Ref. .

Our findings should, on the other hand, not be restricted to the particular neuron model studied here. We expect that stochastic resonance may be introduced to nearly any threshold system when the full dynamics under periodic forcing are reduced to a renewal process by resetting the forcing to a noise-independent phase after each threshold crossing. Periodic forcing may then be recovered by adjusting the noise such that threshold crossings occur around the predefined reset phase, whence periodic forcing is recovered, and optimal output attained. Unless the fixed reset phase has a counterpart in the physical system under study, this resonance is obviously an artefact of a simplification carried too far.

###### Acknowledgements.

We would like to thank G. T. Einevoll for critically reading an earlier version of the manuscript and two anonymous referees for helpful comments. HEP is supported by an EU Marie-Curie-Fellowship.