# insight review articles

Exploring complex networks
Steven H. Strogatz
Department of Theoretical and Applied Mechanics and Center for Applied Mathematics, 212 Kimball Hall, Cornell University, Ithaca, New York 14853-1503, USA (e-mail: strogatz@cornell.edu)

The study of networks pervades all of science, from neurobiology to statistical physics. The most basic issues are structural: how does one characterize the wiring diagram of a food web or the Internet or the metabolic network of the bacterium Escherichia coli? Are there any unifying principles underlying their topology? From the perspective of nonlinear dynamics, we would also like to understand how an enormous network of interacting dynamical systems — be they neurons, power stations or lasers — will behave collectively, given their individual dynamics and coupling architecture. Researchers are only now beginning to unravel the structure and dynamics of complex networks.

268

N

But networks are inherently difficult to understand, as the following list of possible complications illustrates. 1. Structural complexity: the wiring diagram could be an intricate tangle (Fig. 1). 2. Network evolution: the wiring diagram could change over time. On the World-Wide Web, pages and links are created and lost every minute. 3. Connection diversity: the links between nodes could have different weights, directions and signs. Synapses in
Box 1 Nonlinear dynamics: terminology and concepts97

Dynamical systems can often be modelled by differential equations dx/dt v(x), where x(t) (x1(t), …, xn(t)) is a vector of state variables, t is time, and v(x) (v1(x), …, vn(x)) is a vector of functions that encode the dynamics. For example, in a chemical reaction, the state variables represent concentrations. The differential equations represent the kinetic rate laws, which usually involve nonlinear functions of the concentrations. Such nonlinear equations are typically impossible to solve analytically, but one can gain qualitative insight by imagining an abstract n-dimensional state space with axes x1, …, xn. As the system evolves, x(t) flows through state space, guided by the ‘velocity’ field dx/dt v(x) like a speck carried along in a steady, viscous fluid. Suppose x(t) eventually comes to rest at some point x*. Then the velocity must be zero there, so we call x* a fixed point. It corresponds to an equilibrium state of the physical system being modelled. If all small disturbances away from x* damp out, x* is called a stable fixed point — it acts as an attractor for states in its vicinity. Another long-term possibility is that x(t) flows towards a closed loop and eventually circulates around it forever. Such a loop is called a limit cycle. It represents a self-sustained oscillation of the physical system. A third possibility is that x(t) might settle onto a strange attractor, a set of states on which it wanders forever, never stopping or repeating. Such erratic, aperiodic motion is considered chaotic if two nearby states flow away from each other exponentially fast. Long-term prediction is impossible in a real chaotic system because of this exponential amplification of small uncertainties or measurement errors
NATURE | VOL 410 | 8 MARCH 2001 | www.nature.com

geometrically regular ways. Meta-complication: the various complications can influence each other. nonlinear differential equations. Food web of Little Rock Lake. We usually hope the laser will settle down to a stable state. Colours indicate different types of interactions: black. (Figure provided by N. A portion of the molecular interaction map for the regulatory network that controls the mammalian cell cycle6. the state of each node can vary in time in complicated ways. These evolve according to four coupled. the connection between them is strengthened. this is the basis of memory and learning. 230 kV.com E2F Plk1 © 2001 Macmillan Magazines Ltd 269 . The biochemical network that controls cell division in mammals consists of a bewildering variety of substrates and enzymes6. because a perfectly coherent array of N lasers would Figure 1 Wiring diagrams for complex networks. inhibitory or excitatory. When coupled neurons fire together repeatedly. a. enzyme actions. and omnivory (feeding on more than one trophic level) is shown by different coloured links to consumers. 5. the present layout of a power grid depends on how it has grown over the years — a case where network evolution (2) affects topology (1). For instance. currently the largest food web in the primary literature5. To make progress. Pink dashed lines are transformers. For example. 6. Martinez). self-synchronization would be the most desirable outcome. with permission. 765 kV and 500 kV. New York State electric power grid. Thorp and H. D. 6. Node diversity: there could be many different kinds of nodes. and the phase and amplitude of its electric field.nature. or beat each other into incoherence? From a technological standpoint. green. Line thickness and colour indicate the voltage level: red. 138 kV and below. Figure provided by K. red. covalent modifications and gene expression. in nonlinear dynamics we have tended to favour simple.insight review articles the nervous system can be strong or weak. In a gene network or a Josephson junction array. Wang). The lines connecting them are transmission lines and transformers. green. stimulations and inhibitions. 345 kV. nearly identical dynamical systems coupled together in simple. Nodes are functionally distinct ‘trophic species’ containing all taxa that share the same set of predators and prey. only a few of which are shown in Fig. brown. b. Height indicates trophic level with mostly phytoplankton at the bottom and fishes at the top. Kohn. polarization. each laser is characterized by its time-dependent gain. grey. (Reproduced from Fig. (Figure provided by J. Now suppose that many identical lasers are arranged side by side in a regular chain20 or ring21. interacting with their neighbours by evanescent coupling or by overlap of their electric fields22. 6a in ref. different fields have suppressed certain complications while highlighting others. Laser arrays provide a concrete example19–24. In the single-mode approximation. blue. 4. These simplifications allow us to sidestep any issues of structural complexity and to concentrate instead on the system’s potentially formidable dynamics.) c a b C14 P48 CAK Transcription P C42 cdk1 cdk Cyclin box P C43 p16 C12 C10 p19ARF PT286 SL1 P P C14 C24 DMP1 C11 C11a cdc25A P C20 C35 C31 Myc:Max CycH Cdk7 C11b C8 P C9 p16 PCNA R11 C1 cycD C34 C22 C3 cdk4/6 C32 Fos Replication R11 PCNA RPA P P CycE R2 CycA Gadd45 R6 p53 R10 S9 CycD p21/Gadd45 CycA P44 P43 P p21 p27 p57 PT380 C7 R2 C21 p68 p68 primase Pol α C23 PY15 P R2 C13 C19 C2 P CycA Skp1 R1 CBF3 cycE PS76 C26 C26 p68 RPA C28 Skp2 R2 C27 C25 C4 cdk2 E20 P PS216 C38 C39 C37 C36 C40 C18 C29 C2 cycA S9 C30 C41 C5 C15 p53 box P Chk1 C-TAK1 C11b DMP1 p19ARF p53 cycB C42 C41 cdc25C PT14Y15 PT161 SL1 P26 Mdm2 DP1 pRb E2F ? APC C41 P C37 C17 P cdk1 C16 Cks1 C6 Wee1 14-3-3 Myt1 P42 cyclins. Will the lasers lock their phases together spontaneously. binding interactions and stoichiometric conversions. Generators and substations are shown as small blue bars. or break up into a standing wave pattern. c. Wisconsin. Dynamical complexity: the nodes could be nonlinear dynamical systems. but periodic pulsations and even chaotic intensity fluctuations can occur in some cases19. pol α NATURE | VOL 410 | 8 MARCH 2001 | www. corresponding to steady emission of light. 1c. Here nodal dynamics (4) affect connection weights (3). Furthermore we usually assume that the network architecture is static. Cannibalism is shown with self-loops.