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20
Jorge Carballido-Landeira
Bruno Escribano Editors
Biological Systems:
Nonlinear Dynamics
Approach
Se MA
SEMA SIMAI Springer Series
Volume 20
Editor-in-Chief
Luca Formaggia, MOX–Department of Mathematics, Politecnico di Milano,
Milano, Italy
Pablo Pedregal, ETSI Industriales, University of Castilla–La Mancha, Ciudad Real,
Spain
Series Editors
Mats G. Larson, Department of Mathematics, Umeå University, Umeå, Sweden
Tere Martínez-Seara Alonso, Departament de Matemàtiques, Universitat
Politècnica de Catalunya, Barcelona, Spain
Carlos Parés, Facultad de Ciencias, Universidad de Málaga, Málaga, Spain
Lorenzo Pareschi, Dipartimento di Matematica e Informatica, Università degli Studi
di Ferrara, Ferrara, Italy
Andrea Tosin, Dipartimento di Scienze Matematiche “G. L. Lagrange”, Politecnico
di Torino, Torino, Italy
Elena Vázquez-Cendón, Departamento de Matemática Aplicada, Universidade de
Santiago de Compostela, A Coruña, Spain
Jorge P. Zubelli, Instituto de Matemática Pura e Aplicada, Rio de Janeiro, Brazil
Paolo Zunino, Dipartimento di Matematica, Politecnico di Milano, Milano, Italy
As of 2013, the SIMAI Springer Series opens to SEMA in order to publish a joint
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THE SERIES IS INDEXED IN SCOPUS
More information about this series at http://www.springer.com/series/10532

Jorge Carballido-Landeira Bruno Escribano
•
Editors
Biological Systems:
Nonlinear Dynamics
Approach
123
Editors
Jorge Carballido-Landeira Bruno Escribano
Department of Physics Department of Modelling and Simulations
University of Oviedo in Life and Materials Sciences
Oviedo, Spain Basque Center for Applied Mathematics
Bilbao, Spain
ISSN 2199-3041 ISSN 2199-305X (electronic)

SEMA SIMAI Springer Series
ISBN 978-3-030-16584-0 ISBN 978-3-030-16585-7 (eBook)
https://doi.org/10.1007/978-3-030-16585-7
© Springer Nature Switzerland AG 2019
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Preface
This book brings together recent achievements in the field of biological systems
from a nonlinear dynamics perspective. During the compilation of this book, we
have received contributions of recent results obtained by the group of international
scientists who attended the 2nd BCAM Workshop on Nonlinear Dynamics in
Biological Systems, held at the Basque Center for Applied Mathematics, Bilbao in
September 2016. These contributions embrace diverse disciplines and use multi-
disciplinary approaches-including theoretical concepts, simulations and experi-
ments-that emphasize the nonlinear nature of biological systems in order to be able
to reproduce their complex behavior. The results included in this book represent
recent progress and not necessarily what was presented at the conference.
The topics included in the book relate to medical applications as well as more
fundamental questions in biochemistry. One such question is the ability to control
chemically driven reaction–diffusion systems that lead to periodic patterns, which
are often observed in biological problems. Here, the authors present a study of this
question as a mathematical control problem, taking into account time-delay feed-
back for both standing waves and travelling waves.
Closer to medical applications, we include a study about the prevention and
treatment of heart diseases such as tachycardia, ventricular fibrillation and arte-
riosclerosis. In particular, this problem is discussed from the perspective of a simple
dynamical model that describes electrical wave propagation through the heart tis-
sue. This approach uses a reaction–diffusion model to simulate the contraction
of the electrical pulse as it propagates through cardiac tissue, a phenomenon that
can lead to tachycardia and fibrillation.
Another important topic with medical applications is neural network growth.
The work presented here involved the performance of a longitudinal graph
theory-based study of in vitro neuronal networks, comparing results from both
simulations and experiments. This is an example of self-organization through an
optimization process that leads to a random small-world network with positive
degree–degree correlations. The phenomenon can be qualitatively described by a
spatial network growth model based on random growth and neuronal migration.
v
vi Preface
This book also includes recent results in fundamental biochemical questions

such as chiral symmetry breaking in polymers, including a brief description of the
mechanical forces at play and a feasible dynamical model for enantioselective
polymerization in closed systems. The presented model accounts for spontaneous
mirror symmetry breaking as a consequence of competing recycling processes, also
in the absence of chiral or mutual inhibition.
Lastly, we present the application of mechanochemical pattern formation in the
cytoplasm. This problem is studied using viscoelastic models for solid and fluid
cases and in the particular case of cardiac cells.
The editors and the authors wish to express their gratitude to the Basque
Government (Eusko Jaurlaritza) for their financial support in the organization of the
2nd BCAM Workshop on Nonlinear Dynamics in Biological Systems. We would
like to thank the Basque Center of Applied Mathematics (BCAM) for their valuable
assistance in logistics, administrative duties and creation of a good atmosphere for
knowledge exchange. We are also thankful to the Berlin Center for Studies of
Complex Chemical Systems (BCSCCS) for providing further economic support.
Oviedo, Spain Jorge Carballido-Landeira

Bilbao, Spain Bruno Escribano
September 2018
Contents
Time-Delay Feedback Control of an Oscillatory Medium . . . . . . . . . . . . 1

Michael Stich and Carsten Beta
Electrophysiological Effects of Small Conductance Ca2 +-Activated K+
Channels in Atrial Myocytes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19
Angelina Peñaranda, Inma R. Cantalapiedra, Enrique Alvarez-Lacalle
and Blas Echebarria
Spontaneous Mirror Symmetry Breaking from Recycling
in Enantioselective Polymerization . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 39
David Hochberg, Celia Blanco and Michael Stich
Self-organized Cultured Neuronal Networks: Longitudinal Analysis
and Modeling of the Underlying Network Structure . . . . . . . . . . . . . . . . 59
Daniel de Santos-Sierra, Inmaculada Leyva, Juan Antonio Almendral,
Stefano Boccaletti and Irene Sendiña-Nadal
Onset of Mechanochemical Pattern Formation in Poroviscoelastic
Models of Active Cytoplasm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 87
Sergio Alonso
vii
About the Editors
Jorge Carballido-Landeira received his Ph.D. degree in Physics from the

University of Santiago de Compostela (Spain) in 2011, where he worked on non-
linear physical dynamics emerging in active confined chemical systems. He is
currently an Assistant Professor at the Department of Applied Physics at the
University of Oviedo, Spain. His research focuses on a multidisciplinary field that
includes nonlinear dynamics, self-organization processes, colloidal and polymer
science, environmental constraints and fluid mechanics in order to gain an under-
standing of complex biological systems.
Bruno Escribano has been a researcher at the Basque Center for Applied
Mathematics since 2014. He holds a Ph.D. in Physics and Mathematics from the
University of Granada, Spain (2010). His highly multidisciplinary research focuses
on such diverse topics as astrophysics, quasi-crystals, biomineralization and pattern
formation in geophysics. His current research interests are related to the modeling
and simulation of biological systems using nonlinear dynamical methods.
ix
Time-Delay Feedback Control
of an Oscillatory Medium
Michael Stich and Carsten Beta
Abstract The supercritical Hopf bifurcation is one of the simplest ways in which a
stationary state of a nonlinear system can undergo a transition to stable self-sustained
oscillations. At the bifurcation point, a small-amplitude limit cycle is born, which
already at onset displays a finite frequency. If we consider a reaction-diffusion system
that undergoes a supercritical Hopf bifurcation, its dynamics is described by the
complex Ginzburg-Landau equation (CGLE). Here, we study such a system in the
parameter regime where the CGLE shows spatio-temporal chaos. We review a type
of time-delay feedback methods which is suitable to suppress chaos and replace it by
other spatio-temporal solutions such as uniform oscillations, plane waves, standing
waves, and the stationary state.
1 Introduction
1.1 Oscillatory Reaction-Diffusion Systems

and Spatio-Temporal Chaos
The spontaneous emergence of patterns is a fascinating phenomenon observed in

many physical, chemical, and biological systems far from thermal equilibrium. Such
patterns may show complex temporal or spatio-temporal dynamics, including chaotic
behavior. Since these patterns are created by the internal dynamics of the system, this
process is called spatio-temporal self-organization and is referred to as pattern forma-
M. Stich (B)
Non-linearity and Complexity Research Group, System Analytics Research Institute,
School of Engineering and Applied Science, Aston University, Aston Triangle B4 7ET,
Birmingham, UK
e-mail: m.stich@aston.ac.uk
C. Beta
Institute of Physics and Astronomy, University of Potsdam,
14476 Potsdam, Germany
e-mail: beta@uni-potsdam.de
© Springer Nature Switzerland AG 2019 1

J. Carballido-Landeira and B. Escribano (eds.), Biological Systems:
Nonlinear Dynamics Approach, SEMA SIMAI Springer Series 20,
https://doi.org/10.1007/978-3-030-16585-7_1
2 M. Stich and C. Beta
tion [1–4]. Typical examples are the patterns found in chemical reaction-diffusion
systems [2, 3], hydrodynamic and liquid crystal systems [3, 5], electrochemical
systems [6], semiconductors and gas-discharge systems [7, 8], optical systems [9],
granular matter [10], the heart [11], the central nervous system [12], and many other
biological and ecological systems [13–18]. The complexity and diversity of self-
organizing systems is mathematically reflected by nonlinear equations, which arise
in a natural way when systems with many interacting elements, biological systems,
or chemical reactions are considered. One common framework to describe these
phenomena is provided by the research field called nonlinear dynamics [19].
An important class of pattern-forming systems are reaction-diffusion systems,
where the coupling of nonlinear reaction kinetics with a diffusive transport process
leads to complex dynamical behavior. The most prominent example of a chemical
pattern-forming reaction-diffusion system is the Belousov-Zhabotinsky reaction. It
consists of the oxidation of malonic acid by bromate ions in an acidic medium,
catalyzed by metal ions. Boris Belousov discovered self-sustained oscillations of
this reaction under stirring conditions [20] and Anatol Zhabotinsky and Art Winfree
later reported target, spiral, and scroll waves in the unstirred system [21–23]. Later,
major advances have been achieved with a modification of the Belousov-Zhabotinsky
reaction using microemulsions, which shows inward traveling spiral and target waves,
Turing structures, standing waves, oscillatory clusters, and other patterns [24, 25].
Another example of a well-studied chemical reaction giving rise to complex temporal
and spatio-temporal behavior is the oxidation of carbon monoxide on platinum single
crystal surfaces under low pressure conditions [26, 27]. A special feature of this
reaction is the occurrence of spatio-temporal chaos and its control [28], which we
will comment on later.
The concept and mathematical structure of reaction-diffusion models are quite
general, and therefore such models may also be applied to pattern-forming physical
and biological systems. In the context of physical systems, reaction-diffusion models
are for instance used to describe the dynamics of localized patterns found in gas-
discharge systems [29]. In the field of living systems which show oscillations or
wave phenomena, reaction-diffusion models are for example successfully applied to
explain excitation waves (and their breakdown) in the heart [11, 30] and aggregation
patterns in slime mold colonies [31].
Depending on the local kinetics in the absence of diffusive coupling, differ-
ent types of reaction-diffusion systems can be distinguished, such as multistable,
excitable, or oscillatory systems [32]. Here, we will focus on oscillatory media that
are characterized by temporally periodic dynamics and may give rise to wave trains,
spirals, or concentric wave patterns. The simplest bifurcation in which stable limit
cycle oscillations emerge from a stationary fixed point is the Hopf bifurcation. Close
to the bifurcation point, it is possible to derive a general model, which holds for
all systems undergoing such a bifurcation, regardless of their specific nature. This
universal equation is the complex Ginzburg-Landau equation [33].
The complex Ginzburg-Landau equation has the interesting property that it dis-
plays not only the aforementioned regular wave patterns, but also different states of
spatio-temporal chaos. In particular, a state of phase turbulence has been described,
Time-Delay Feedback Control of an Oscillatory Medium 3
where oscillation phases behave chaotic while amplitudes are still regular, and a
state of defect (or amplitude) chaos, where phases and amplitudes are displaying
turbulent dynamics. Along with other properties of this equation, these dynamical
states have been characterized in detail [34]. The complex Ginzburg-Landau equation
(CGLE) [33, 35] reads
∂t A = (1 − iω)A − (1 + iα)|A|2 A + (1 + iβ)∇ 2 A, (1)
where A is the complex oscillation amplitude, ω the linear frequency parameter, α

the nonlinear frequency parameter, β the linear dispersion coefficient, and ∇ 2 the
Laplacian operator. While the equation contains a term with the parameter ω, it could
be scaled out with a transformation A → A exp(−ωt) to leave the model with two
parameters only. However, we prefer to keep this additional parameter in the model
because in this way we keep the relation to a real reaction-diffusion system as ω
scales with the distance μ to the Hopf bifurcation point: ω ∝ μ−1 .
1.2 Time-Delay Feedback: Global and Local Control

Schemes
Among the most challenging questions in this field is the control of chaotic behavior.
Chaos control has evolved into a rapidly expanding domain of research in its own
right [36]. Inspired by the work of Ott, Grebogi, and Yorke (OGY method) [37],
chaos control was first realized for low-dimensional systems. While the OGY method
becomes too laborious when implemented in high-dimensional systems, empirical
control schemes were designed that can be readily applied to spatially extended
systems. The most widely known variant of these schemes was proposed by Pyragas
in his seminal paper in 1992 [38] that has been modified and extended in various
ways (see e.g. [39]). By generating a control signal from the difference between the
actual system state and a time-delayed one, powerful methods can be designed to
influence the dynamics of a nonlinear system in a subtle, self-generated way. Different
approaches based on delay differential equations, nonlinear dynamics, and control
theory meet in this interdisciplinary research field [36, 40]. Using such methods,
control of spacetime chaos has been studied for a number of different systems such
as optical devices [41], lasers [42], and chemical systems [28]. A large body of work
on the control of spatio-temporal chaos has been performed with the catalytic CO
oxidation on Pt(110), a surface catalytic reaction that has served as a model system
to investigate pattern formation in reaction-diffusion systems. Space-time chaos in
the CO oxidation is characterized by the statistics of topological phase defects [43,
44], and it can be suppressed by global time-delayed feedback [28, 45, 46] as well as
periodic forcing [47, 48]. Also exotic variants of mode-dependent feedback have been
considered [49]. Moreover, the laser-induced generation of localized wave sources
has been used to suppress spatio-temporal chaos in the CO oxidation system [50–52].
In general, we may distinguish global and local feedback schemes. In the first
case, the same feedback force is uniformly applied to the system at each point in
space. In the case of a local scheme, the feedback acts in a spatially resolved fashion
and may take different values at different points in space. Local and global feed-
back schemes can be seen as limiting cases of a more general situation, where the
control signal contains both local and global contributions. Such intermediate cases
are found in many experimental systems, in particular in the biomedical context. A
prominent example is spreading depolarization in the cerebral cortex during migraine
and stroke. In an effort to model this system, both local and nonlocal feedback has
been considered [53, 54]. A second widely studied example is the control of chaotic
states in excitable media that underlie cardiac arrhythmias [55]. Global control (con-
ventional defibrillation) as well as spatially resolved techniques are studied in this
field; for an overview see [56]. Also in other systems, combinations of global and
local feedback terms may appear. A prototypical system, in which delayed feed-
back methods have been studied in great detail, is charge transport in semiconductor
devices [7]. Here, global feedback typically arises from the global nature of the volt-
age drop across the device, whereas the space-dependent interface charge density
allows for the implementation of either global or local feedback terms [57–60].
Also in the context of the CGLE, both global and local feedback schemes of dif-
ferent kinds have been studied with the aim of controlling native chaotic states. The
CGLE under the effect of global time-delay feedback has been studied by Battogtokh
et al. [61, 62]. A global feedback scheme of Pyragas type (time-delay autosynchro-
nization – TDAS) was considered by Beta and Mikhailov [63]. The effect of local
feedback on the CGLE was first investigated by Socolar and coworkers [64, 65].
Later, Silber and coworkers have extended the study of local feedback in the CGLE
to a generalized form of the Pyragas scheme that includes spatial shifts in addition
to time delay to stabilize traveling wave solutions [66, 67]. Here, we will discuss
the CGLE for a one-dimensional medium with a combination of local and global
time-delayed feedback. It has been introduced in Ref. [68] and reads
∂t A = (1 − iω)A − (1 + iα)|A|2 A + (1 + iβ)∂x x A + F, (2)

F = μeiξ m l (A(x, t − τ ) − A(x, t)) + m g ( Ā(t − τ ) − Ā(t)) , (3)
where L
1
Ā(t) = A(x, t) dx (4)
L 0
denotes the spatial average of A over a one-dimensional medium of length L. The

parameter μ describes the feedback strength and ξ characterizes a phase shift between
the feedback and the current dynamics of the system. The parameters m g and m l
denote the global and local contributions to the feedback, respectively. If m l = 0,
the case of global time-delayed feedback is retrieved, which has been studied in
Ref. [63]. We adopt the notation used in Ref. [68], which slightly differs from the
one in Ref. [63]. Obviously, if m g = 0, then we have a case of purely local feedback.
In the simulations shown below, we typically choose m g + m l = 1 constant, thus

reducing the number of parameters by one. If μ = 0, the model reduces to the standard
complex Ginzburg-Landau equation. In the following we will review our research
on this model [68–72].
2 Uniform Solutions
We anticipate that the system introduced in Eq. (2) shows a range of regular and irreg-
ular solutions, and we start our discussion with the simplest ones, namely uniform
oscillations and the uniform stationary state.
2.1 Existence of Uniform Oscillations
The uniform periodic solution (“uniform oscillations”) is given by
A0 (t) = ρ0 e−iΩt . (5)
It is a solution of Eq. (2) with the amplitude and frequency given by

ρ0 = 1 + μ(m g + m l )χ1 , (6)
Ω = ω + α + μ(m g + m l )(αχ1 − χ2 ) . (7)
Here, χ1,2 denote effective modulation terms that can be positive or negative. They
arise from the feedback and hence depend on ξ and τ ,
χ1 = cos(ξ + Ωτ ) − cos ξ, (8)

χ2 = sin(ξ + Ωτ ) − sin ξ. (9)
In general, no explicit analytic solution for Eqs. (6) and (7) can be given because χ1,2
also depend on Ω. Nevertheless, the solution can be computed using root-finding
algorithms, as done in [69].
For μ = 0, the solution reduces to ρ0 = 1 and Ω = ω + α which is the well-
known uniform oscillatory solution of the standard CGLE. It is well-known for the
latter equation that if 1 + αβ < 0 (the Benjamin-Feir-Newell criterion), the homoge-
neous periodic solution A0 = e−i(ω+α)t is unstable and a regime of spatio-temporal
chaos is observed. Below, we will consider the case where this criterion is fulfilled.
Solving Eqs. (6) and (7) as a function of τ for all other parameters fixed, one
observes that depending on the value of the feedback strength μ, there may be
multiple solutions for an interval of τ . This is shown in Fig. 1 where the frequency
(a) and amplitude (b) of oscillations are shown for a set of parameter for which the
1.2 1.6
(a) 1.4 (b)
1.1
1.2
1
1
T 0.9 0.8
0.6
0.8
0.4
0.7
0.2
0.6 0
0.6 0.7 0.8 0.9 1 1.1 1.2 0 0.2 0.4 0.6 0.8 1 1.2
τ τ
Fig. 1 a The period of oscillations T = 2π/Ω as determined by Eq. (7) as a function of τ for
μ = 0.3 (dashed), μ = 0.66 (solid), μ = 1.0 (dotted-dashed). b Periods T = 2π/Ω (solid) and
amplitudes ρ0 (dashed, from Eq. (6)) as a function of τ are shown for μ = 1.5. For more information
see text. The other parameters are α = −1.4, ω = 2π − α (hence the unperturbed period is equal
to 1), ξ = π/2. Figure from Ref. [71]
period of oscillations without feedback is equal to unity. If the feedback strength is

small and we vary τ , the oscillation frequency is not constant, but always relatively
close to the value of the frequency of oscillations for μ = 0. However, if the value
of μ is larger than a critical threshold, there is an interval of τ for which the Eq. (7)
has three solutions. If an even larger value of μ = 0 is chosen, the interval of τ in
which multiplicity of solutions occurs becomes larger. In Ref. [71] we showed that
as τ becomes larger, also the multiplicity areas become larger.
2.2 Stability of Uniform Oscillations
Above we have stated that there are parameter areas with multiplicity of oscillatory
solutions. If we are interested in the stability of such solutions, we have to take
this multiplicity into account. Nevertheless, the treatment of the stability analysis is
quite general, so there is no need to introduce the solution until a given numerical
experiment.
To perform a linear stability analysis of uniform oscillations with respect to spatio-
temporal perturbations, we express the complex oscillation amplitude A as the super-
position of a homogeneous mode H with spatially inhomogeneous perturbations,
A(x, t) = H (t) + A+ (t)eiκ x + A− (t)e−iκ x . (10)
Inserting Eq. (10) into Eq. (2), and assuming that the amplitudes A± are small, we
obtain
∂t H = (1 − iω)H − (1 + iα)|H |2 H + μ(m l + m g )eiξ (H (t − τ ) − H (t)), (11)

∂t A+ = (1 − iω)A+ − (1 + iα)(2|H |2 A+ + H 2 A∗− ) − (1 + iβ)κ 2 A+
+μm l eiξ (A+ (t − τ ) − A+ ), (12)
∂t A∗− = (1 + iω)A∗− − (1 − iα)(2|H |2 A∗− + H ∗2 A+ ) − (1 − iβ)κ 2 A∗−
+μm l e−iξ (A∗− (t − τ ) − A∗− ). (13)
See Appendix A of Ref. [69] for details of this derivation. The solution of Eq. (11)
is given by uniform oscillations H (t) = H exp(−iΩ t) identical to Eq. (6), i.e.,
H = ρ and Ω = Ω. The equations for A+ and A∗− include terms proportional to
m l . They represent the local contributions to the feedback term and constitute the
difference between Eqs. (12), (13) and Eqs. (7)–(9) of Ref. [63]. To investigate linear
stability of uniform oscillations with respect to spatio-temporal perturbations, we
make the ansatz
A+ = A0+ exp(−iΩt) exp(λt), (14)

A∗− = A∗0
− exp(iΩt) exp(λt), (15)
where λ = λ1 + iλ2 is a complex eigenvalue. The sign of its real part determines
stability. After substituting Eqs. (14), (15) into Eqs. (12) and (13) we arrive at the
following eigenvalue equation:
F = (A + iB − iλ2 + D1 + iD2 )(A − iB − iλ2 + C1 + iC2 ), (16)
where we have defined
F = (1 + α 2 )ρ 4 , (17)
A = 1 − λ1 − 2ρ − κ , 2 2
(18)
B = Ω − ω − 2αρ 2 − βκ 2 , (19)
C1 = μm l e−λ1 τ cos(ξ + Ωτ + λ2 τ ) − μm l cos ξ, (20)
C2 = −μm l e−λ1 τ sin(ξ + Ωτ + λ2 τ ) + μm l sin ξ, (21)
D1 = μm l e−λ1 τ cos(ξ + Ωτ − λ2 τ ) − μm l cos ξ, (22)
D2 = μm l e−λ1 τ sin(ξ + Ωτ − λ2 τ ) − μm l sin ξ. (23)
For details of this derivation the reader is referred to Appendix B of Ref. [69]. This
system of equations can then be evaluated to check for the stability of the oscillatory
solution. As usual, the criterion for the first instability is that the real part of the
largest eigenvalue crosses zero while all others are still negative. When this happens,
the imaginary part reveals whether the instability itself is of oscillatory nature. At
the same time, the wavenumber of the most unstable perturbation tells us whether
the instability is associated with a spatial periodicity.
In extensive investigations performed in Ref. [69], a variety of stability diagrams

were shown and discussed and examples of different instabilities were presented.
As the main result it was found that there is a wide range of possible delay times τ
such that as the feedback strength μ is increased, the chaotic state is indeed replaced
by a uniform oscillatory state. For nonzero μ, the feedback-induced solution is dif-
ferent from the uniform oscillation without feedback and hence the magnitude of
the feedback term is nonzero as the feedback-induced solution is stabilized. This is
characteristic for a so-called invasive control [68].
It is worth mentioning that the stability analysis captures not the transition from
the fully chaotic state to unifrom oscillations. Rather, it describes the first instabil-
ity encountered from the uniform oscillatory state upon a decrease of the feedback
strength. While instabilities of different kinds (imaginary parts nonzero, wavenum-
bers zero, etc.) are possible, arguably the most common instability is the one with
vanishing imaginary part and nonzero wavenumber. This corresponds to the onset of
a pattern with spatial periodicity but with no additional temporal component besides
the underlying uniform oscillations. Such a pattern describes standing waves, as
shown in Fig. 2 and investigated in more detail in Sect. 3.1. Figure 2 also shows for
comparison space-time plots for uniform oscillations and developed defect (ampli-
tude) chaos.
(a) (b) (c)
Fig. 2 Main spatio-temporal solutions: a uniform oscillations, b standing waves, d spatio-temporal

chaos. Shown are space-time diagrams in gray scale for |A| (top panels) and ReA (bottom panels)
for a time interval of t = 25 in the asymptotic regime and system size L = 256. The delay time
is τ = 0.5 and the values of μ are μ = 0.50 (a), μ = 0.15 (b), μ = 0 (c). Black (white) denotes
low (high) values of the respective quantity (rescaled for each simulation). For |A|, these values
are (|A|min , |A|max ) = (0.94, 1.13) (b), (|A|min , |A|max ) = (0.15, 1.2) (c). For (a), the amplitude
is constant |A| = 1.085. The other parameters are as in Fig. 1, and β = 2. These patterns were first
reported in Ref. [68]
2.3 Existence and Stability of the Stationary State
The CGLE has been derived to describe the onset of oscillations in a Hopf bifurcation
from a stationary state (fixed point). In the standard version of the CGLE that we
utilize, the stationary state is simply represented by a vanishing amplitude |A| =
ρ ≡ 0 (the phase is no longer defined). This trivial solution exists always, so we only
have to investigate its stability.
Following the approach used in Ref. [69], we use the ansatz (10) and perform a lin-
ear stability analysis with respect to spatio-temporal perturbations. After separating
uniform and periodic modes, we obtain
∂t H = (1 − iω)H + μ(m l + m g )eiξ (H (t − τ ) − H (t)), (24)

∂t A+ = (1 − iω)A+ − (1 + iβ)κ 2 A+ + μm l eiξ (A+ (t − τ ) − A+ ), (25)
∂t A∗− = (1 + iω)A∗− − (1 − iβ)κ 2 A∗− + μm l e−iξ (A∗− (t − τ ) − A∗− ), (26)
where as usual higher order terms in H and A± have been neglected. These equations
are decoupled and can be studied independently. In order to explore the linear stability
of the state H = 0 with respect to uniform perturbations, we set
H = H 0 exp(λt), (27)
with H 0 an initial amplitude and λ a complex eigenvalue. Inserting Eq. (27) into
Eq. (24), yields the following characteristic equation
λ = 1 − iω + μ(m l + m g )eiξ (e−λτ − 1). (28)
Separating real and imaginary parts, Eq. (28) can be written in the form

λ1 = 1 + μ(m l + m g ) e−λ1 τ cos(ξ − λ2 τ ) − cos ξ , (29)

λ2 = −ω + μ(m l + m g ) e−λ1 τ sin(ξ − λ2 τ ) − sin ξ . (30)
These equations can be solved numerically, searching for λ1 = 0 and determining

the parameter area where the stationary state is stable. Some analytic operations
can be made using the Lambert W function [73]. In the literature on control of
oscillations, the stabilization of this stationary state is also called amplitude death,
since oscillations are no longer observed [74]. In particular, Eq. (28) is equivalent to
the amplitude death condition for a single Hopf oscillator.
3 Spatio-Temporal Solutions
In this section, we consider patterns that show a space-dependence. In particular,

we study standing wave patterns observed in a large parameter region between uni-
form oscillations and turbulence. Then, we give a short overview over other patterns
observed in this system.
3.1 Standing Waves
One of the ways uniform oscillations can become unstable is when spatially periodic
perturbations can increase and give rise to standing waves with wavelength 2π/kc .
We follow Ref. [70] to describe the derivation of the solution of standing waves. To
be more precise, at onset we have
λ1 (kc ) = 0 , ∂k λ1 (kc ) = 0 , and λ2 (kc ) = 0 , with kc = 0,
where λ is the leading eigenvalue of the stability analysis of uniform oscillations, and
λ1 and λ2 its real and imaginary part. The ansatz to capture such patterns analytically
is therefore
A(x, t) = H (t) + Bk (t)(eikx + e−ikx ), (31)
with H being the uniform mode and Bk the complex amplitude of the mode with
wavenumber k. This ansatz can also be interpreted as a rightgoing wave (with
wavenumber k) and a leftgoing one (with wavenumber −k) sharing the same ampli-
tude and thus describing a standing wave pattern.
Before inserting ansatz (31) into Eq. (2), let us express several terms appearing in
Eq. (2) for the case that A(x, t) is given by Eq. (31), of particular relevance being
|A|2 A = |H |2 H + 4|Bk |2 H + 2H ∗ Bk2

+ (eikx + e−ikx )(H 2 Bk∗ + 2|H |2 Bk + 3|Bk |2 Bk )
+ (e2ikx + e−2ikx )(2H |Bk |2 + H ∗ Bk2 )
+ (e3ikx + e−3ikx )|Bk |2 Bk , (32)
and
Fl = μm l eiξ [(H (t − τ ) − H (t))

+ (eikx + e−ikx )(Bk (t − τ ) − Bk (t))], (33)
Fg = μm g eiξ (H (t − τ ) − H (t)). (34)
Equation (33) assumes that the space-dependent part of A(x, t) does not contribute
to the average Ā(t). This is fulfilled if L → ∞ or L = 2π n/k (with n = 1, 2, ...). In
the following, we will neglect the higher-order harmonic contributions in Eq. (32),
which close to onset is well-justified. Inserting finally ansatz (31) into Eq. (2), and
using Eqs. (32)–(34), we obtain
dH d Bk
+ (eikx + e−ikx ) = (1 − iω)(H + (eikx + e−ikx )Bk )
dt dt
−(1 + iα)(|H |2 H + 4|Bk |2 H + 2H ∗ Bk2
+(eikx + e−ikx )(H 2 Bk∗ + 2|H |2 Bk + 3|Bk |2 Bk ))
−(1 + iβ)k 2 (eikx + e−ikx )Bk
+μ(m l + m g )eiξ (H (t − τ ) − H (t))
+μm l eiξ (Bk (t − τ ) − Bk (t))(eikx + e−ikx ). (35)
Now, we separate the equation into space-dependent and space-independent parts

and arrive at
dH
= (1 − iω)H − (1 + iα)(|H |2 H + 4|Bk |2 H + 2Bk2 H ∗ )
dt
+μ(m l + m g )eiξ (H (t − τ ) − H (t)), (36)
d Bk
= (1 − iω)Bk − (1 + iα)(H 2 Bk∗ + 2|H |2 Bk + 3|Bk |2 Bk )
dt
−(1 + iβ)k 2 Bk + μm l eiξ (Bk (t − τ ) − Bk (t)). (37)
In the case of Bk = 0, the first equation reduces to the equation describing the mode of
the homogeneous periodic solution. However, the coupling terms in the first equation
of Eq. (36) tell us that if Bk = 0, the uniform mode H is different from the uniform
oscillations (5) and we cannot use Eq. (6).
To solve Eq. (36), we assume the solution to be of the following form [75]:
H = H0 e−iΩ0 t , (38)
−i(Ω0 t+γ )
Bk = Bk0 e . (39)
Thus, both modes oscillate at the same frequency Ω0 , while there is a phase shift γ
between the modes. The real amplitudes are given by H0 and Bk0 , respectively.
After inserting (38), (39) into (36), we obtain
2 −2i(Ω0 t+γ ) 2iΩ0 t
(−iΩ0 )H = (1 − iω)H − (1 + iα)(H02 + 4Bk02
+ 2Bk0 e e )H
iξ iΩ0 τ
+μ(m l + m g )e (e − 1)H, (40)
(−iΩ0 )Bk = (1 − iω)Bk − (1 + iα)(H02 e−2iΩ0 t e2i(Ω0 t+γ ) + 2H02 + 3Bk0
2
)Bk
iξ iΩ0 τ
−(1 + iβ)k Bk + μm l e (e
2
− 1)Bk . (41)
Assuming H nonzero in the first equation, and Bk in the second, this simplifies to
2 −2iγ
0 = 1 + iΩ0 − iω − (1 + iα)(H02 + 4Bk0
2
+ 2Bk0 e )
+μ(m l + m g )eiξ (eiΩ0 τ − 1), (42)
0 = 1 + iΩ0 − iω − (1 + iα)(H02 e2iγ + 2H02 + 3Bk0
2
)
iξ iΩ0 τ
−(1 + iβ)k + μm l e (e
2
− 1). (43)
These equations can be separated into real and imaginary parts. We write them as
0 = 1 − (H02 + 2Bk0
2
(2 + cos 2γ )) − 2α Bk0
2
sin 2γ + μ(m l + m g )χ1s , (44)
0 = Ω0 − ω − α(H0 + 2Bk0 (2 + cos 2γ )) + 2Bk0 sin 2γ + μ(m l + m g )χ2 , (45)
2 2 2 s
0 = 1 − k 2 − (H02 (2 + cos 2γ ) + 3Bk0

2
) + α H02 sin 2γ + μm l χ1s , (46)
0 = Ω0 − ω − βk 2 − α(H02 (2 + cos 2γ ) + 3Bk0 2
) − H02 sin 2γ + μm l χ2s , (47)
where χ1,2
s
are given by
χ1s = cos(ξ + Ω0 τ ) − cos ξ, (48)

χ2s = sin(ξ + Ω0 τ ) − sin ξ, (49)
For a given k, Eqs. (44)–(47) can be solved numerically through root-finding algo-
rithms, giving solutions for H0 , Bk0 , Ω0 , and γ . There, the wavenumber k is provided
by the eigenvalue problem studied in [69], where we use either kc (at threshold) or
kmax (away from threshold). Combining Eqs. (31), (38), (39), the family of standing
wave solutions can be written as
A SW = e−iΩ0 t (H0 + 2Bk0 cos(kx)e−iγ ). (50)
This solution is plotted in Fig. 3 for an example set of parameters together with
the result of a simulation of the CGLE for the same parameters. The agreement is
striking, given that the mode separation involves several approximations (neglecting
higher-order harmonics, assuming that the space-dependent part of A(x, t) does not
contribute to the average Ā(t), and that the wavenumber is determined by the linear
stability analysis of uniform oscillations).
3.2 Other Patterns
The CGLE admits a range of different spatio-temporal solutions, so it should not

come as a surprise that also the system in the chaotic regime subjected to a time-
delay feedback is able to show the stabilization of multiple solutions with simulta-
neous space and time dependence. Simulations revealed, among others: (a) traveling
waves that arise as standing waves undergo an instability to translational motion or
Fig. 3 Amplitude profiles 1

for μ = 0.7 (other
parameters as in Fig. 2).
0.8
Shown are a simulation
oscillation amplitudes
(solid black curve) and the
theoretical result using the 0.6
theoretically obtained wave
number (dashed blue curve).
We display only a part of the 0.4
medium (total size,
L = 128). This solution was
0.2
first presented in Ref. [70] full simulation
analytical solution
0
50 60 70 80 90 100
space
breathing; (b) plane waves, i.e. waves with a constant amplitude; (c) traveling phase
flips on a background of standing waves; and (d) bound states of pacemakers [68].
A theoretical description of these patterns is still lacking.
4 Summary and Discussion
Control of chaotic states has been investigated intensively in the last decades within
the realm of nonlinear dynamics, control theory, and dynamical systems theory,
among others fields. In this chapter, we have reviewed a particular case of spatio-
temporal chaos and its control, namely the amplitude chaotic state in the complex
Ginzburg-Landau equation (CGLE) and its control via a time-delay feedback term.
The latter has global and local contributions and the terms are constructed in the
spirit of the Pyragas idea of a feedback that vanishes in the case of control.
As the CGLE describes a system with underlying small-amplitude, sinusoidal
oscillations, the main solution that can be induced by the feedback are uniform oscil-
lations. We have described the corresponding solution analytically. As the controlled
solution is different than the solution without feedback, the control is invasive, i.e.,
the control force does not vanish as the solution is stabilized. The solution without
feedback is unstable and not found in the simulations.
Between the uniform oscillations and the turbulent state, standing waves are found.
The corresponding solution has been described analytically and the onset of the stand-
ing waves has been confirmed to be an instability of the uniform oscillations with
respect to spatial perturbations. Again, the feedback is invasive, and the standing
wave solution is not a solution of the CGLE without feedback. Other, more com-
plicated patterns like traveling waves, phase flips and bound states have been found
numerically but an analytic description has not been performed yet.
One pattern that in the case of control has vanishing feedback is the steady state.
This noninvasive state is also called amplitude death, as the oscillations disappear
completely. It can be interpreted as time-delay feedback pushing the system below

the supercritical Hopf bifurcation. Together with the homogeneous oscillations it is
one of the two uniform patterns studied. For such solutions, the global and local
feedback terms become identical. However, as we have seen, this does not imply
that global and local terms act equally with respect to stability of these solutions.
The description of standing waves as uniform oscillations that become unstable with
respect to perturbations of a certain wavelength is an example of this.
The validity of the existence and stability criteria of the above-mentioned patterns
is limited to one spatial dimension. While the existence criteria for the uniform
patterns generalize into two dimensions, the stability analysis will be fundamentally
different as the allowed perturbations can depend on both dimensions simultaneously.
A study of this situation would be an interesting subject of future work.
Another limitation of the work presented in this chapter is that the fundamental
CGLE parameters were chosen to correspond to spatio-temporal chaos and are not
systematically varied. While we believe that we have now understood central parts
of the process of controlling amplitude chaos in the CGLE via time-delay feedback,
the same need not to be true for phase chaos or other fundamental solutions of the
CGLE. Relevant questions to be asked include how the time-delay feedback – global,
local, or mixed – could change the stability area of uniform oscillations, which are
stable for the unperturbed CGLE.
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Electrophysiological Effects of Small
Conductance Ca2+ -Activated K+
Channels in Atrial Myocytes
Angelina Peñaranda, Inma R. Cantalapiedra, Enrique Alvarez-Lacalle

and Blas Echebarria
Abstract Atrial fibrillation (AF), a cardiac arrhythmia characterized by an abnormal

heart rythm originated in the atria, is one of the most prevalent cardiac diseases.
Although it may have diverse causes, genetic screening has shown that a percentage of
pacients suffering of AF present a genetic variant related to disregulation of calcium-
activated potassium (SK) channels. In this paper we review the main characteristics of
these channels and use several mathematical models of human atrial cardiomyocytes
to study their influence in the form of the atrial action potential. We show that an
overexpression of SK channels results in decreased action potential duration and,
under some circumstances, it may give rise to alternans, suggesting a pro-arrhythmic
role of this current. This effect becomes more important at higher pacing rates.
Nevertheless, we also find it to protect against spontaneous calcium release induced
afterdepolarizations, acting in this case as an antiarrhythmic factor.
1 Introduction
In the heart, contraction is driven by an electrical wave that propagates through the
atria and ventricles. This wave has its origin in the ion fluxes that cross the car-
diomyocytes’ cell membrane, producing a transient change in electrical polarity of
the membrane, known as an action potential, that propagates from cell to cell. The
cardiac action potential is the result of a complex interplay of several transmembrane
A. Peñaranda · I. R. Cantalapiedra · E. Alvarez-Lacalle · B. Echebarria (B)

Departament de Física, Universitat Politècnica de Catalunya,
08028 Barcelona, Spain
e-mail: angelina.penaranda@upc.edu
I. R. Cantalapiedra
e-mail: inmaculada.rodriguez@upc.edu
E. Alvarez-Lacalle
e-mail: enric.alvarez@upc.edu
B. Echebarria
e-mail: blas.echebarria@upc.edu
© Springer Nature Switzerland AG 2019 19

J. Carballido-Landeira and B. Escribano (eds.), Biological Systems:
Nonlinear Dynamics Approach, SEMA SIMAI Springer Series 20,
https://doi.org/10.1007/978-3-030-16585-7_2
20 A. Peñaranda et al.
currents. Sodium, potassium and calcium ions are involved in determining the appro-
priate shape of the action potential. Sodium flow into the cell is responsible for the
initial depolarization current, while potassium and calcium ions move in and out
afterwards, balancing each other in order to sustain the depolarization during the
duration of a typical action potential (AP).
In the cell, most calcium ions are kept in a store, known as the sarcoplasmic
reticulum (SR). When calcium enters the cell (due, for example, to an increase in
transmembrane potential), it binds to some calcium-activated receptors at the surface
of the SR (the ryanodine receptors, RyR), that open, releasing the content of the SR
in a proccess known as calcium induced calcium release (CICR). After relaxation,
calcium is reuptaken into the SR and brought out of the cell by the Na–Ca exchanger.
Thus, the duration of the action potential is closely linked to the calcium transient
and dysfunctions in one affect the other. In effect, under anomalous conditions,
calcium release from the SR can occur spontaneously, resulting in an increase of
transmembrane voltage (through the action of the Ca–Na exchanger), that may give
rise to a depolarization, making it a possible source of ectopic heartbeats and cardiac
arrhythmias.
In recent years, there has been increased attention into another feedback mecha-
nism between calcium transients and transmembrane potential, in this case mediated
by small-conductance Ca2+ -activated potassium channels, or SK channels [31, 37,
50]. These channels belong to a large family of Ca2+ -activated potassium channels,
that also include big (BK) and intermediate (IK) conductances (see Sect. 2.1 for more
details). The three types of channels differ in their pharmacological properties. SK
channels, for instance, and contrary to BK or IK channels, are blocked by apamin, a
neurotoxin found in bee venom.
SK channels were first discovered in skeletal muscle [3], and subsequently they
have been found in a wide range of cell types: neuronal, endothelia, epithelia, etc.
[40, 49]. In the heart, the first evidence of the presence of SK channels and a Ca2+
activated K + current (I K Ca ) was found in a rat ventricular cell line [45], but these
channels have been found in a wide variety of species and types, from rabbit to rat
and human [34, 48], and in the pulmonary vein, ventricle, atria and in pacemakers
[7, 8]. SK channels have been found to be preferentially expressed in atria compared
to the ventricles [47].
In the atria, there is ample evidence that the SK current contributes to action poten-
tial repolarization [43, 47]. Inhibition of SK currents has been shown to prolong AP
duration (APD) in mouse and human atrial myocytes [39, 47]. Similarly, ablation of
SK channels resulted in a significant prolongation of APD [25] while overexpres-
sion of SK showed a significant APD shortening [51]. Following their role in the
atrial action potential, SK channels have been recognized to play an important role in
atrial fibrillation. The relation between expression of SK and AF has been confirmed
by genome-wide association studies, revealing that polymorphisms in the calcium-
dependent potassium-channel gene KCNN3 are an important risk factor for atrial
fibrillation (AF) [14, 33, 35]. This result suggests that SK channels may represent a
potential therapeutic target for the treatment of atrial arrhythmias.
Electrophysiological Effects of Small Conductance … 21
SK channels expression may also play a role in atrial remodelling, i.e., changes in
size, shape, structure, and function of the atria occurring in patients with AF. The first
work to point in this direction was by Ozgen et al. [34] where fast pacing increased the
presence of SK channels, as manifested in the APD abbreviation and the sensitivity of
the current to apamin. Other works link this remodelling with enhanced trafficking
of SK channels to the membrane during rapid atrial tachycardia [36]. However,
SK channel expression has been observed to be significantly reduced in right atrial
appendages recovered from chronic AF patients [48] and in patients with permanent
AF [26]. A possible explanation for these contradictory results proposes that AF may
result in the up-regulation of SK channels as the initial response but, with progression
of the disease, they might be down-regulated [50].
Despite the evidence relating disfunctions in SK channels with AF, their pro-
or anti-arrythmic role is not clear. For instance, a decrease in the SK current in
a SK knock-out mouse model resulted in action potential prolongation and atrial
fibrillation [25]. Similar results have been observed in the canine left atrium [21]. In
these studies the prolongation of APD was accompanied by increased occurrences
of early after depolarization (EAD), increased APD heterogeneity, occurrences of
electrical alternans, and wave breaks. This protective role of SK current against AF
agrees with the observation that SK current was decreased in human atria tissue in
patients with AF [39]. These results, however, seem to be in contradiction with the
observation that inhibition of SK channels significantly reduced AF inducibility [12,
13, 35, 38] and SK channel overexpression resulted in predisposition to induced atrial
arrhythmias [51]. Most probably, these contradictory results stem from the various
possible mechanisms leading to atrial arrythmias and AF. Since SK current helps
repolarization it is expected to protect against afterdepolarization related arrhythmias,
while its role in decreasing the APD may help initiation and maintainence of reentries.
In this regard, little has been done regarding modeling the effect of SK chan-
nels in cardiac electrophysiology in order to unveil different pro- or anti-arrhythmic
effects of inhibition of SK channels. Recently, Kennedy et al. [22] studied the effect
of calcium-sensitive potassium currents on voltage and calcium alternans in a ven-
tricular cell model but no similar study has been performed for atrial cell models.
In this paper we will perform a first analysis of its effect on atrial action potential
models. We review existing models of I K Ca currents (not necessarily in cardiac cells)
and then study the effect of this current in the cardiac atrial action potential. Due
to the undeterminancy in the details of some features of the SK current, we will
study its effects under different values for its half-activation calcium concentration,
its maximal conductance, and the calcium binding cooperativity. This should be the
starting point in order to test whether SK channels could avoid or limit the effect of
spontaneous calcium release (SCR) on the generation of afterdepolarizations.
Another random document with
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A pretty good thing, and I suppose there’s nursemaids and
governesses for all the children, too, hey?
Maxwell
Why, as a matter of fact there are—but not exactly in the way
you mean. You see, they run a sort of a boarding school, too, the
people that run this model tenement do, and they take care of all the
children there—keep ’em night and day, feed ’em and dress ’em and
teach ’em trades and all that sort of thing. They let them come to see
you on Sunday, but you’re relieved of all responsibility and your wife
of all the trouble.
Ryan
I see. But what’s to become of my job when I’m going off to this
here model tenement to sleep nights? Don’t you know that a janitor
has to sleep in the building he’s taking care of?
Maxwell
Well, you’re not in love with your job, are you? You’d be willing to
do some other work where there was a chance of advancement and
better pay, wouldn’t you?
Ryan
Sure I would. Where am I going to get it?
Maxwell
Why, as to that, I guess Mr. Morris or I could get you a job
somewhere. You’re a big husky fellow and pretty steady, I know. I
could get you a job in the shipping department of a factory I am
interested in, if you wanted it.
Ryan
What would the hours be?
Maxwell (laughing)
Why, you’re going pretty fast, Ryan. I don’t know just what the
hours would be yet, but I suppose they’d be from half past eight to
six or something like that. That’s easier than your hours now, isn’t it?
Ryan
Much easier. Now, what has all this got to do with my wife having
too many children?
Maxwell
Why, these people think that Mrs. Ryan has had too many
children. Some of these people are doctors, and they say it’s bad for
a woman’s health to have so many children.
Ryan
I see. Well, what would you like to have me do about it?
Maxwell
About the model tenement?
Ryan
No, about my wife having any more children.
Maxwell
Well, you know—you see I’m not—what they think is— Oh,
damn it, I don’t know about this part of the thing, Ryan. It’s out of my
line. I’m not a philanthropist. I’m just talking for these people
because they know I know you. About that question you’ll have to
talk to Mrs. Dannenberg or Mr. Morris.
Ryan
Mr. Morris—he’s the little guy that was standing on a chair when
I came in to-night, ain’t he?
Maxwell
Yes.
Ryan
Is he a doctor?
Maxwell
No, I don’t think he is.
Ryan
He came into my place the other day selling some sort of a little
doctor book.
Maxwell
I know; he does that because he’s a philanthropist.
Ryan
Do people make a living by being philanthropists?
Maxwell
No—well, as a matter of fact, many people do make a pretty fat
living out of it, but Mr. Morris doesn’t. He’s a sculptor—a man that
makes statues, you know.
Ryan
Oh, I know what a sculptor is, all right. What statues did Mr.
Morris make?
Maxwell
Why, he made that statue over in the corner, for one thing.
(Ryan goes over and examines the purple statue.)
Ryan
Do you like this statue, Mr. Johnson?
Maxwell (laughing)
I’m not a judge of such things, Ryan. But people who know about
art say that it’s very good indeed.
Ryan
Is it a good likeness?
Maxwell
It’s not supposed to be a portrait; it’s a sort of a fancy statue. It
represents the “Emancipation of Woman”—woman freed, you know.
Ryan
Freed from what?
Maxwell
Why, from overwork, and excessive child-bearing and all that
sort of thing, you know.
Ryan
Well, that lady’s freed from child-bearing, all right. She’s perfectly
safe on that score. (He goes back to his chair and pours out another
drink.) But now let’s get back to your friends’ proposition. I’m to get a
day job and come home nights—is that right?
Maxwell
That’s right.
Ryan
The children are taken off my wife’s hands, and she doesn’t have
to do any more cooking or washing or anything, hey?
Maxwell
That’s it, Ryan.
Ryan
And the place we’d live would be pretty much like this apartment,
would it?
Maxwell
Just as comfortable, at any rate.
Ryan
My wife would have no work to do; I’d work in the daytime and
come home nights—there’d be no kids to take care of—it would be
pretty much like the life that you and your wife have, wouldn’t it, Mr.
Johnson?
Maxwell
Ah—why, yes, Ryan, pretty much the same. What do you think of
the proposition?
Ryan
Well, I’ll tell you. Of course I wouldn’t like to have the children go
away—though they are a lot of trouble—but I suppose they’d be
better taken care of than we could do, so, if that was all there was to
it, I’d say go ahead. And it’d be all right for me, too, if I was a
bachelor. But it’s on account of my wife that I feel I’ve got to say,
“Excuse me!”
Maxwell
Why, you don’t begrudge your wife a little rest, do you, Ryan?
Ryan
Would I begrudge the old woman a little rest? Yes, by God, I
would, if a little rest meant having nothing to do all day except sit
around and talk to her friends and run around town. It’s just on that
account that I say nix to your whole proposition. Now you think I’m a
slave driver, I suppose. Well, I ain’t, Mr. Johnson, but I’ve lived with a
woman thirteen years, and what I’m telling you I didn’t get out of no
books—it’s facts!
Listen, Mr. Johnson. What you want us to do is to live just the
sort of life that you and your wife live—no children to take care of, no
washing nor cooking nor nothing. Well, what I say is, excuse me!
That may be all right for you and your wife—she don’t need no
housework nor children nor nothing to keep her busy. Her time is full
of all sorts of useful things—I know that. (Maxwell stirs uneasily and
looks at the floor.) But with Annie, my wife, it’s a different proposition
altogether. She’s one of them women—and there’s a lot more of
them than you think—that can’t stand living in a nice regular
apartment with nothing to do. I know because we tried it when we
was first married. It’s like what the old fellow said: “Satan finds some
mischief still for idle hands to do”!
Maxwell
True enough! And—
Ryan
Listen, Mr. Johnson. When we was first married, Annie was one
of these here idle hands they tell of. She was an idle hand for three
years, and Satan, as the old fellow said, certainly found some
mischief for her to do. Not anything real bad—there’s no real harm in
Annie—but it was mischief all right. For the first three years we was
living in a nice apartment in Brooklyn. I wasn’t a janitor, then; I was
driving a truck. I was out all day and I got home every night at six or
seven o’clock. All Annie has to do is to get my breakfast and supper
and keep the place clean. Does she do it? Sure she does, for the
first four or five weeks! Then she gets to making friends with other
women in the building and going out to matinees and vaudeville and
all that sort of thing. That’s all right—I can afford it—I don’t care if
she has a good time; but then what does she do? I give her two
dollars in the morning to go out and buy a good supper for me when
I come home. I come home and she ain’t in yet, and the lady in the
next flat gives me the groceries that she’s sent home. And what is
they? A little chipped beef and a box of Saratoga chips and some
baker’s bread. About fifty-cent’s worth. When she gets home I ask
her where she’s been. Why, Mrs. Eindorfer has took her to a
spiritualist meeting, and she’s spent the rest of that money to look
into a glass ball or have her fortune told or some such foolishness.
Now this goes on for nearly three years. It ain’t all spiritualists’
meetings; it’s all sorts of things. She makes all sorts of friends,
women and men, too; I had to beat a couple of ’em up. The flat
wasn’t kept up; I run into debt; my meals wasn’t cooked right or on
time, and Annie was half sick all the time just from running around
entertaining herself. I ain’t blaming her. She wasn’t to blame. And
what was to blame? The apartment house was to blame. When
Peter was born, after we’d been married three years, and I gave up
trucking and moved out of that apartment house and got a job as
janitor, everything was all right. And everything’s been all right ever
since.
Maxwell (thoughtfully)
And the apartment house was to blame?
Ryan
Believe me, Mr. Johnson, the only part of an apartment house to
live in is the basement, where you can have a regular home. I been
a janitor for ten years, and I seen these apartment houses do queer
things to families. They don’t seem to have no children when they
live in apartment houses, that’s one thing. And there ain’t no coal to
bring up and the washing goes out, and there ain’t nothing for them
to do but just make fools of themselves. And sometimes there’s a
good many divorces been caused by these here apartment houses.
And there’d be a good many more divorces if a lot of husbands knew
what went on when they was downtown at business.
Understand me, Mr. Johnson. I don’t mean you and your wife at
all. You ain’t that sort of people, but what I do say is for my wife, and
for a lot of women with more education and more money than she’s
got, the only sort of life is doing housework and taking care of
children all day long. So Annie and me will stay down in the
basement, much obliged to you, unless we go out of New York to live
in a little house in the country sometime. And Annie’ll have just as
much work to do there. She’s one of them women that wasn’t meant
to be idle. And now I guess I’ll go downstairs.
(He rises and goes toward the door. Maxwell sits silent for a
moment and then rises a little unsteadily. He holds out his hand to
Ryan for a second and then drops it and starts.)
Maxwell
Ryan, I—er—why, I guess you’re right, after all. I’ll tell my friends
what you said.
Ryan
All right. No hard feelings, I hope.
Maxwell
Not at all; that’s all right. Good night, Ryan.
Ryan
Good night, Mr. Johnson.
(He goes out.)
Maxwell
There’s a man that’s master in his own home, at any rate. (He
lights a cigar and walks around the room with his hands in his
trousers pockets, coming to a halt in front of the purple statue. He
looks at it reflectively.) Satan finds some mischief still—(A pause)—
for idle hands to do. For idle hands to do. For idle hands to do. For
idle hands—
(The doorbell is rung violently. Maxwell starts and runs out into
the hall. He returns with Helen, who is very much out of breath. As
she runs into the room the combs drop from her hair, which falls over
her face and shoulders. She throws herself on the chaise-longue.
Maxwell sits beside her and tries to push her hair back from her
face.)
Nellie! What’s the matter?
(Helen sobs without answering.)
Helen
It’s those nasty Martins and that nasty policeman and that nasty
Lionel Morris.
(The bell rings again. Maxwell goes to the door and admits
Lionel.)
Lionel
Oh, I’m awfully glad you got back all right, Miss White. I jumped
into a taxi as soon as that brute of a policeman came, and then I met
all the rest of the crowd at the studio and everybody said, “Where’s
Miss White?” So I came right up here to find out if you’d got home.
Maxwell
For God’s sake, will somebody tell me what’s happened?
Lionel (sitting on the pianola bench)

Why, you see—
Helen
Be still. You see, Max, we were all at the Mortons’ studio, and
Adrian Wolfe made a speech about those nasty striking miners in
California or wherever it is that everybody is wearing mourning for
and parading and all that and this—and Mr. Morris said: “Let’s walk
up to Union Square and hold an open air meeting to protest.” So we
went up there and I made a speech and there was a crowd and I
saw a policeman there, but I thought it would be all right, and then
Mr. Morris made a speech and he said something about trampling on
a bloodstained flag, and the policeman told him to stop, and he
called the policeman a myrmidon, and some more policemen came
and broke up the meeting, and he ran away and wouldn’t help me;
and I ran down into the subway, and I don’t see how he dares show
his face in here!
Maxwell
I’ll talk to him presently, but quiet down a little. You’d better go in
your room and fix up your hair.
(Helen rises to go out. She stops in the doorway and turns to
Maxwell.)
Helen
I won’t go to that nasty Amaranth this summer, Maxwell.
Maxwell
No, you won’t go to Amaranth.
Helen
Then will you get an automobile?
Maxwell
No, I won’t get an automobile.
Helen
Then what—
Maxwell
I am going to take that two thousand dollars and buy, with the
assistance of the building and loan association, a small house in a
city called Joplin, in the State of Missouri. It will not be a large house,
but I think that you will not find the time hanging heavy on your
hands. My brother has a wholesale grocery there, and I dare say he
will take me into the business, especially as I have a little money to
invest. And I’ll come home to luncheon every day. Missouri is a fertile
State. My brother has six children.
Lionel
But, Miss White—Mr. Johnson!
(Helen goes down the hall to her room. Maxwell walks up to
Lionel.)
Maxwell
My wife’s name is not Miss White but Mrs. Johnson—Mrs.
Maxwell Johnson, of Joplin, Missouri. Get that? Do you know what
keeps me from dropping you down the elevator shaft?
Lionel
What—what do you mean?
Maxwell
It’s the janitor. Yes, Ryan, the fellow down in the basement with
nine children that you and Mrs. What’s-her-name wanted to
segregate. He told me all about you to-night. You’re nothing but a by-
product! The apartment house is the real devil in this pretty little play
—the apartment house is responsible for Feminism and Socialism
and Anarchism and Eugenics and pups like you. You’re just a sort of
bad substitute for the movies—that’s all you are. The apartment
house breeds the whole bunch of you—the apartment house and its
artificial, lazy, good-for-nothing life.
(Lionel starts toward the door hurriedly, but stops as if shot
when the telephone bell rings close to his ear. He comes back into
the room and Maxwell goes to the telephone.)
Maxwell
Hello!... What’s that?... Yes, this is Mr. Johnson.... No, I don’t
think so. Hold the wire and I’ll see. (With his hand over the
transmitter he looks into the room.) Nellie!
Helen (coming into the living room with her hair

in a long braid, wearing a blue tea gown)
Yes, Max?
Maxwell
What is your name?
Helen
Why, Helen, of course, stupid.
Maxwell
Helen what?
Helen
Helen Johnson.
Maxwell
Not Miss Helen White?
Helen
No! No! No!
Maxwell (smiling)
Well, that’s all right, then. There’s a cop down-stairs with a
warrant for the arrest of a Miss Helen White and a Mr. Lionel Morris,
charged with making incendiary speeches in Union Square. They
think that Morris’s taxicab stopped at this building, and the policeman
is going through all the apartments. He’ll be here in a minute. (In the
receiver.) All right, Sam, it’s all right. Thanks for tipping me off.
(Maxwell reënters the room and sits on the pianola bench. Helen
reclines, with some dignity, on the chaise-longue. Lionel crouches
behind the purple statue.)
Maxwell (meditatively)
Satan finds some mischief still—
Helen
What are you saying, Max?
Maxwell
Oh, I was just thinking of the janitor. I had quite a talk with him
after you left.
(The doorbell rings, and Maxwell admits a large policeman.)
Policeman
Excuse me, sir; it’s just a matter of form. I’m looking for a couple
of them Anarchist-Suffrage-I. W. W. bugs. It’s just a matter of form.
The man’s name is Lionel Morris and the woman’s name is Helen
White. Are you Lionel Morris?
Maxwell
No; my name is Maxwell Johnson. The janitor knows me, and so
do a lot of people in the building.
Policeman
Thank you, sir. It’s just a matter of form. Now, madam—it’s just a
matter of form—are you Helen White?
Helen
No, I am not Helen White. I am Mrs. Maxwell Johnson.
Policeman
Thank you, madam, thank you; it’s just a matter of form. You see
these parties is incendiaries; they called me a mermaiden. Now, just
two more questions—it’s just a matter of form: Is Miss Helen White
here?
Helen
No, Helen White is not here.
Maxwell
And I’m glad she isn’t here, officer.
Policeman
You may well be that, sir; you may well be that. Now, is Lionel
Morris here?
(There is a pause, during which the purple statue shakes
slightly.)
Maxwell
Well, what do you think about it, officer? Take a look around the
place. Want to look in the dumb waiter or down the kitchen sink?
Policeman
Oh, I know he’s not here, Mr. Johnson, and I’m sorry to have
troubled you. Much obliged to you. Good night, sir.
Maxwell
Have a drink before you go?
Policeman
Well, I hadn’t ought to, but I guess I will, thanks.
(Helen pours the whiskey and Maxwell and the policeman lift
their glasses.)
Policeman
Well, here’s how, sir.
Maxwell
Here’s Joplin!
Helen
Oh, I’ll drink that.
(She takes a sip from Maxwell’s glass.)
Policeman
What’s that, something new?
Maxwell
No, it’s old as Adam and Eve.
Policeman
Well, it’s a new one on me. Thank you, sir. Good night.
Maxwell
Good night. (The policeman goes out. After the door slams shut,
Lionel stands up, but remains behind the purple statue.) Hadn’t you
better go to some other apartment house? The cop’s gone down the
elevator. He’ll be gone by the time you get downstairs. (Lionel goes
out and as he turns he brushes against the purple statue, which
topples on its pedestal. He bangs the door shut after him violently,
and the statue falls to the floor and breaks into several pieces.
Maxwell and Helen look at it for a moment and then turn to each
other and laugh.)
Helen
Oh, look what’s happened to the “Emancipation of Woman”!
CURTAIN
Transcriber’s Notes
Punctuation, hyphenation, and spelling were made
consistent when a predominant preference was found in the
original book; otherwise they were not changed.
Simple typographical errors were corrected; unbalanced
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obvious, and otherwise left unbalanced.
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Biological Systems Nonlinear

Dynamics Approach Jorge

Nonlinear Dynamics of Discrete and Continuous Systems

IUTAM Symposium on Exploiting Nonlinear Dynamics for

Nonlinear Circuits and Systems with Memristors:

Nonlinear Dynamics of Structures Systems and Devices

An Introduction to Complex Systems Society Ecology and

Electric Aircraft Dynamics: A Systems Engineering

A Practical Approach to Fracture Mechanics Jorge Luis

Global Nonlinear Dynamics for Engineering Design and

THE SERIES IS INDEXED IN SCOPUS

More information about this series at http://www.springer.com/series/10532

ISSN 2199-3041 ISSN 2199-305X (electronic)

This book also includes recent results in fundamental biochemical questions

Oviedo, Spain Jorge Carballido-Landeira

Time-Delay Feedback Control of an Oscillatory Medium . . . . . . . . . . . . 1

Jorge Carballido-Landeira received his Ph.D. degree in Physics from the

Michael Stich and Carsten Beta

1.1 Oscillatory Reaction-Diffusion Systems

The spontaneous emergence of patterns is a fascinating phenomenon observed in

© Springer Nature Switzerland AG 2019 1

∂t A = (1 − iω)A − (1 + iα)|A|2 A + (1 + iβ)∇ 2 A, (1)

where A is the complex oscillation amplitude, ω the linear frequency parameter, α

1.2 Time-Delay Feedback: Global and Local Control

∂t A = (1 − iω)A − (1 + iα)|A|2 A + (1 + iβ)∂x x A + F, (2)

denotes the spatial average of A over a one-dimensional medium of length L. The

In the simulations shown below, we typically choose m g + m l = 1 constant, thus

2.1 Existence of Uniform Oscillations

The uniform periodic solution (“uniform oscillations”) is given by

A0 (t) = ρ0 e−iΩt . (5)

It is a solution of Eq. (2) with the amplitude and frequency given by

χ1 = cos(ξ + Ωτ ) − cos ξ, (8)

period of oscillations without feedback is equal to unity. If the feedback strength is

2.2 Stability of Uniform Oscillations

A(x, t) = H (t) + A+ (t)eiκ x + A− (t)e−iκ x . (10)

∂t H = (1 − iω)H − (1 + iα)|H |2 H + μ(m l + m g )eiξ (H (t − τ ) − H (t)), (11)

A+ = A0+ exp(−iΩt) exp(λt), (14)

F = (A + iB − iλ2 + D1 + iD2 )(A − iB − iλ2 + C1 + iC2 ), (16)

where we have defined

In extensive investigations performed in Ref. [69], a variety of stability diagrams

(a) (b) (c)

Fig. 2 Main spatio-temporal solutions: a uniform oscillations, b standing waves, d spatio-temporal

2.3 Existence and Stability of the Stationary State

∂t H = (1 − iω)H + μ(m l + m g )eiξ (H (t − τ ) − H (t)), (24)

λ = 1 − iω + μ(m l + m g )eiξ (e−λτ − 1). (28)

These equations can be solved numerically, searching for λ1 = 0 and determining

In this section, we consider patterns that show a space-dependence. In particular,

3.1 Standing Waves

λ1 (kc ) = 0 , ∂k λ1 (kc ) = 0 , and λ2 (kc ) = 0 , with kc = 0,

|A|2 A = |H |2 H + 4|Bk |2 H + 2H ∗ Bk2

Fl = μm l eiξ [(H (t − τ ) − H (t))

Now, we separate the equation into space-dependent and space-independent parts

0 = 1 − k 2 − (H02 (2 + cos 2γ ) + 3Bk0

χ1s = cos(ξ + Ω0 τ ) − cos ξ, (48)

A SW = e−iΩ0 t (H0 + 2Bk0 cos(kx)e−iγ ). (50)

3.2 Other Patterns

The CGLE admits a range of different spatio-temporal solutions, so it should not

Fig. 3 Amplitude profiles 1

λ1 (kc ) = 0 , ∂k λ1 (kc ) = 0 , and λ2 (kc ) = 0 , with kc = 0,