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NL3516 Catalytic hypercycle 1

NL3516 Catalytic hypercycle


The concept of the hypercycle has been invented in the nineteen seventies
in order to characterize a functional entity that integrates several autocatalytic
elements into an organized unit (Eigen, 1971; Eigen & Schuster, 1977, 1978a,b). A
catalytic hypercycle is defined as a cyclic network of autocatalytic reactions (figure 1).
Autocatalysts, in general, compete when they are supported by the same source of
energy or material. Hypercyclic coupling introduces mutual dependence of elements
and suppresses competition. Consequently, the fate of all members of a hypercycle is
identical with that of the entire system and, in other words, no element of a hypercycle
dies out provided the hypercycle as such survives. The current view of biological
evolution distinguishes periods of dominating Darwinian evolution based on variation,
competition, and selection interrupted by rather short epochs of radical innovations
often called major transitions (Maynard Smith & Szathmary, 1995; Schuster, 1996).
In the course of biological evolution major transitions introduce higher hierarchical
levels. Examples are: (i) the origin of translation from nucleic acid sequences into
proteins including the invention of the genetic code, (ii) the transition from independent
replicating molecules to chromosomes and genomes, (iii) the transition from the
prokaryotic to the eukaryotic cell, (iv) the transition from independent unicellular
individuals to differentiated multicellular organisms, (v) the transition from solitary
animals to animal societies, and (vi) presumably a series of successive transitions from
animal societies to man. All major transitions introduce a previously unknown kind
of cooperation into biology. The hypercycle is one of very few mechanisms that can
deal with cooperation of otherwise competing individuals. It is used as a model system
in prebiotic chemistry, in evolutionary biology, in theoretical economics as well as in
cultural sciences.
The simplest example of a catalytic hypercycle is the elementary hypercycle. It is
described by the dynamical system1
n
!
dxi X
= xi fi xi1 fj xj1 xj ; i, j = 1, 2, . . . , n; i, j = mod n . (1)
dt j=1

The catalytic interactions within a hypercycle form a directed closed loop comprising
all elements, often called Hamiltonian arc: 1 2 3 . . . n 1 (figure 1).
Hypercycles are special cases of replicator equations of the class
n n X n
!
dxi X X
= xi aij xj ajk xj xk ; i, j, k = 1, 2, . . . , n (2)
dt j=1 j=1 k=1

with aij = fi i1,j ; i, j = mod n.1 For positive rate parameters and initial conditions
inside the positive orthant2 the trajectory of a hypercycle remains within this orthant:
1
The mod n function implies a cyclic progression of integers, 1, . . . , n 1, n, 1, . . . . The symbol i,j
represents Kroneckers symbol: = 1 for i = j and = 0 for i 6= j.
2
The notion of orthant refers to the entire section of a Cartesian coordinate system in which the signs
of variables dont change. In n-dimensions the positive orthant is defined by {xi > 0 i = 1, 2, . . . , n}.
NL3516 Catalytic hypercycle 2

Figure 1. Definition of hypercycles. Replicator equations as described by the


differential equation (2) can be symbolized by directed graphs: The individual species
are denoted by nodes and two nodes are connected by an edge, j i, if and only if
aij > 0. The graphs of hypercycles consist of single Hamiltonian arcs as sketched on the
left hand side of the figure. These dynamical systems are permanent independently
of the choice of rate parameters fi . For n 5 they represent the only permanent
systems but for n 6 the existence of a single Hamiltonian arc is only a sufficient
but not a necessary condition for permanence. The graph on the right hand side, for
example, does not contain a Hamiltonian arc but the corresponding replicator equation
is permanent for certain choices of rate parameters (Hofbauer & Sigmund, 1998).

fi > 0, xi (0) > 0 i = 1, 2, . . . , n = xi (t) > 0 t 0. In other words, none of


the variables is going to vanish and hence, the system is permanent in the sense that
no member of a hypercycle dies out in the limit of long times, limt xi (t) 6= 0 i =
1, . . . , n. The existence of a Hamiltonian arc, i.e. a closed loop of directed edges visiting
all nodes once, is a sufficient condition for permanence (Hofbauer & Sigmund, 1998). It
is also a necessary conditions for low-dimensional systems with n 5, but there exist
permanent dynamical systems for n 6 without a Hamiltonian arc; one example is
shown in figure 1.
The dynamics of equation (1) remains qualitatively unchanged when all rate
parameters are set equal: f1 = f2 = . . . = fn = f , which is tantamount to a barycentric
transformation of the differential equation (Hofbauer, 1981). The hypercycle is invariant
with respect to a rotational change of variables, xi = xi+1 with i = 1, 2, . . . , n; imod n,
it has one equilibrium point in the center and its dynamics depends exclusively on n.
Some examples with small n are shown in figure 2. The systems with n 4 converge
towards stable equilibrium points, whereas the trajectories of the equation (1) with
n 5 approach limit cycles. Independently of n, elementary hypercycles do not sustain
chaotic dynamics.
Hypercycles have two inherent instabilities, which are easily illustrated for
molecular species: (i) The members of the cycle may also catalyze the formation of non-
members that do not contribute to the growth of the hypercycle and thus hypercycles are
vulnerable to parasitic exploitation (Eigen & Schuster, 1978a,b), and (ii) concentrations
of individual species in oscillating hypercycles (n 5) go through very small values and
NL3516 Catalytic hypercycle 3

Figure 2. Solution curves of small elementary hypercycles. The figure shows the
solution curves of equation (1) with f1 = f2 = . . . = fn = 1 for n = 2 (upper
left picture), n = 3 (upper right picture), n = 4 (lower left picture), and n = 5
(lower right picture). The initial conditions were: x1 (0) = 1 (n 1) 0.025 and
xk (0) = 0.025 k = 2, 3, . . . , n. The sequence of the curves xk (t) is: k = 1 full black
line, k = 2 full grey line, k = 3 hatched black line, k = 4 hatched grey line, and
k = 5 black line with long hatches. The cases n = 2, 3, and 4 have stable equilibrium
points in the middle of the concentration space c = (1/n, 1/n, . . . , 1/n); equation (1)
with equal rate parameters, n = 4, and linearized around the midpoint c exhibits a
marginally stable center and very slow convergence is caused by the non-linear term
that becomes smaller as the system approaches c. For n = 5 the midpoint c is unstable
and the trajectory converges towards a limit cycle (Hofbauer et al., 1991).

these species might become extinct through random fluctuations. More elaborate kinetic
mechanisms can stabilize the system in case (ii). Exploitation by parasites, case (i),
can be avoided by compartmentalization. Competition between different hypercycles is
characterized by a strong non-linearity in selection (Hofbauer, 2002): Once a hypercycle
has been formed and established, it is very hard to replace it by another hypercycle.
Epochs with hypercyclic dynamics provide explanations for once for ever decisions or
frozen accidents.

See also Biological evolution; Optimization


NL3516 Catalytic hypercycle 4

Further Reading
Eigen, M. 1971. Selforganization of Matter and the evolution of biological
macromolecules. Naturwissenschaften, 58: 465-523
Eigen, M. & Schuster, P. 1977. The hypercycle. A principle of natural self-organization.
Part A: Emergence of the hypercycle. Naturwissenschaften, 64: 541-565
Eigen, M. & Schuster, P. 1978a. The hypercycle. A principle of natural self-organization.
Part B: The abstract hypercycle. Naturwissenschaften, 65: 7-41
Eigen, M. & Schuster, P. 1978b. The hypercycle. A principle of natural self-organization.
Part C: The realistic hypercycle. Naturwissenschaften, 65: 341-369
Hofbauer, J. 1981. On the occurrence of limit cycles in the Volterra-Lotka equation.
Nonlinear Analysis, 5: 1003-1007
Hofbauer, J. 2002. Competitive exclusion of disjoint hypercycles. Zeitschrift f
ur
Physikalische Chemie, 216: 35-39
Hofbauer, J., Mallet-Paret, J. & Smith, H.L. 1991. Stable periodic solutions for the
hypercycle system. Journal of Dynamics and Differential Equations, 3: 423-436
Hofbauer, J. & Sigmund, K. 1998. Evolutionary Games and Replicator Dynamics,
Cambridge, UK: Cambridge University Press
Maynard Smith, J. & Szathmary, E. 1995. The Major Transitions in Evolution, Oxford,
UK: W. H. Freeman & Co.
Schuster, P. 2002. How does complexity arise in evolution? Complexity, 2(1): 22-30

Peter Schuster