Stochastic Resonance addition to BEMS paper Kendra Krueger University of Colorado at Boulder, Electrical Engineering Department October 2011 Kendra.Krueger@gmail.com A New Outlook on Noise Noise has always been seen as a nuisance, the intrinsic harbinger of chaos; unavoidable and inherent of the universe. Every system that lingers away from the ideal model will expectantly be plagued by some sort of random fluctuations which will cloud or degrade the output result. However, this perspective of noise and its properties may be shifting. In the past ten years researchers have begun to observe strange benefits within noisy systems. This phenomenon has come to be known as stochastic resonance. Noise itself is defined to be random background fluctuations, which over time should average to zero. The term stochastic describes a system which behaves randomly with no predictable nature, or is dominated by noisy processes. In most systems when an input signal is combined with noise, the output will also have an increase in noise. However, in stochastic resonance (SR) systems the addition of noise to an input signal will actually improve the resultant output. Dynamical Stochastic Resonance Two models exists to explain this phenomenon, the first is known as dynamical SR which requires a weak periodic input signal and noise that is greater in magnitude than the input (Benzi, Sutera, & Vulpiani, 1981). Now, intuitively we could imagine that adding these two signals together will create a very noisy periodic function at the output, but this, in fact, is not the case. This behavior can be modeled using non-linear bistable equations. Figure 1: Double well potential model of dynamical SR One intuitive way to imagine a bistable system is to think of a particle in a double well, illustrated in figure 1. Here the particle will tend to rest at points of low potential energy. As it does so, the potential energy function is also fluctuating periodically, and is susceptible to random fluctuations. With just the energy provided by the periodic oscillation, the particle will never leave its comfortable position. However, with the additional push from random fluctuations the particle is capable of overcoming the barrier and repositioning itself within the adjacent state, or potential. The behavior of this particle can be modeled using the following equation: Where 𝑥 describes the position of the particle, 𝑈 is the potential of the well, the sin function acts as the periodic oscillation, and ) is the noise. We can envision that the periodic function modulates the depth of the well, making one or the other deeper at any given time. The noise provides the extra push to get the particle over the hill. This combination of functions allows for the particle to be in either state depending on conditions. If we think of the periodic function as being caused by an input signal or voltage that is changing the system or potential we can easily determine the ideal amount of noise needed to reach optimal behavior. Most input/output systems can be evaluated using a signal-to-noise ratio (SNR). In a common intuitive system, as noise is added the SNR will decrease, and the signal will become degraded or destroyed. However, in a SR system the noise will actually improve the SNR. This result was observed in a study which sought to measure the effects of noise on the generations of action potentials in sensory nerves within crayfish tailfins (Douglass, Wilkends, Pantazelou, & Moss, 1993). The SNR obtained matched the behavior seen in figure 2, a fast rise to maximum SNR, and then a steady decrease as noise intensity is increased. This represents the classic SR response in a system, suggesting an optimal noise level. The maximum signal to noise ratio occurs when the noise energy is half the barrier energy [Bulsara 1996]. Figure 2: SNR behavior in SR systems To further evaluate this type of system one can determine a power spectrum analysis. This sort of method examines the critical frequency required to enhance the input signal. This is direct evidence of a resonance phenomenon, a fundamental frequency at which the system will uniquely respond to an external signal. The noise enhancement of the power spectral density by crayfish mechanoreceptors has been measured to be 4.5dB at the optimal noise intensity [Douglass 1993]. Figure 3: Power Spectrum behavior in SR systems Non-Dynamical Stochastic Resonance A model more applicable to biological systems is known as ‘threshold SR’ or ‘non-dynamical SR’. In these systems the noise aids in enhancing a non-detectable signal in order to reach a given detection threshold. For non-dynamical SR, the input signal does not have to be temporally periodic; instead, the frequency of the oscillations may vary in time around an average value (Gingl, Kiss, & Moss, 1995). As the signal + noise combination crosses threshold, an encoded discretized signal is produced. It is believed that encoded within this signal is information about the original data, information which would remain undetectable without the addition of noise. Figure 4: 'Non-dynamic' SR model For experiments which rely on this type of discreet data or threshold detection, information correlation is used as a type of SNR. This information correlation examines the similarity between input and output bits. Mechanoreceptors in rat afferent nerves have shown SR behavior with the non-dynamical model (Collins, Imhoff, & Grigg, 1996). Figure 5: Information Correlation behavior of SR Biochemical Mechanisms of Stochastic Resonance After these initial studies, researchers have attempted to dig deeper to find the biochemical source of stochastic resonance. Ion channels are responsible for regulating the electrochemical equilibrium of the cell membrane by allowing specific types of ions to flow inwards or outwards, creating voltage modifying currents (Kandel, Schwartz, & Jessell, 2000). Each channel is made of a long string of molecules which forms a protein that transcends the lipid bilayers of the membrane, creating a pathway between the extracellular environment and the internal cytoplasm of the cell. These proteins are binary or bistable in nature exhibiting a conformation associated with either open or closed states, no median state exists. Furthermore there are many different activation triggers for each ion channel. Some can be chemically activated, tension activated (such as those in mechanoreceptor cells) or voltage gated. Voltage gated channels have been studied predominantly in the case of stochastic resonance and as such will be discussed further here. It has been observed that the neuronal response to a repeated stimulus is not identical every time (Goychuk & Hanggi, 2000). This calls to attention the probabilistic behavior of the ion channel, which will, in effect, cause noise within the membrane voltage potential. The current generated by ion channels along with the variance can be derived using the following equation In these equations represents the conductance of a single ion channel, N is the number of channels, p(V) is the probability of the channel being open and V is the membrane potential. From these equations the noise can be calculated and is known as the coefficient of variation (CV) Looking at this equation, it can be inferred that the noise should decrease proportional to the square root of the number of channels. This calculation has manifested itself in the broad assumption that at large densities of ion channels, the effective noise can be ignored. This, however, is not the case. When examining the firing threshold in a Hodgkin-Huxley signal neuron model, the probability of firing as a function of input amplitude is not discreet. Instead there is a gradual rise indicating a factor of ‘relative spread’ (RS). Figure 6: The effects of neuronal noise on relative spread of firing probabilities This relative spread can be directly correlated with an increase in channel noise. This suggests that with higher amounts of noise, the probability of a response occurring at low amplitudes is higher than for conditions with low noise. The RS can be calculated using also known as the error function. These results suggest that neuronal noise is more involved in threshold activity than previously thought. Soon new models may need to be modified in order to include the effects of noise. Closing Remarks It is apparent that stochastic resonance has established itself as a legitimate process in which systems are able to amplify a subthreshold signal with the addition of noise. In all of these models, both dynamical and non-dynamical, the noise is visualized as being entrained on the incoming signal, riding atop it and aiding in the transferring of information to the detection mechanism. However, another point of view may be to see the noise as altering the threshold itself. This concept was approached in the channel noise study, but not fully recognized. Examining this phenomenon with this perspective may lead to new insights on how systems develop these sorts of capabilities. This further leads to questions about how systems directly adapt to use this process, or if it is just a byproduct of evolution. One might ask, however, if nature strived to detect subthreshold signals, than why not make the threshold lower? We can’t forget that noise isn’t something new; nature has evolved in the presence of noise and thus sought to benefit from the start. By utilizing the energy that is already present within the background, the system does not need to build a more robust and perhaps less efficient detection system. Nature will always follow the path of least resistance. Counter-intuitively noise, usually thought of as a hindrance or resistance, and may actually provide the more energetically favorable route. A new hypothesis has emerged known as the Living Matter Way (LMW) which theorizes that the complexity of information is key to biological communication, and that noise is the vehicle through which this complex information is transferred (West & Grigolini, 2010). This theory revolves around the production and transmission of 1/f noise, a spectrum of noise discovered by Schottky in electrical devices. This form of noise is inherent in all sorts of process from biological to economic. What arises through this analysis is the idea of high densities of information being encoded in a low energy system. It is interesting to suddenly hear a mention of energy in this sort of context. In all of the preceding experiments the quest was always to determine the relationships of input and output information, but there was no mention as to the energy that may also be transferred in this process. Investigations into this aspect of the process will inevitable conjure up discussion on the entropy rates associated with stochastic resonance amplification and rectification. Stochastic resonance not only sheds light on the complex functioning of the nervous system, but also begins to pave the way for greater understanding of the fundamentals of information transfer and the values of noise in our natural paradigm. Technological developments have historically fought against the probabilistic fatalism of noise, but with inspiration from these newly discovered phenomena, we may in time learn to embrace an inexorably stochastic universe and thrive within the chaos. Works Cited Bahar, S., & Moss, F. (2004). Stochastic resonance and synchronization in the crayfish caudal photo receptors. 188 (pg 81-97). Benzi, R., Sutera, A., & Vulpiani, A. (1981). The mechanism of stochastic resonance. 14 (L453L457). Collins, J. J., Imhoff, T. T., & Grigg, P. (1996). 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The Living Matter Way to exchange information. 75 (pg 475478). White, J. A., Rubinstein, J. T., & Kay, A. R. (2000). Channel noise in neurons. 23 (pg131-137).