Fig. 8. How connectivity influences network dynamics. A, The Lyapunov exponent X is plotted for each of T time steps for different values of the branching parameter O. The branching parameter governs the sum of transmission probabilities from each unit of the network. When O is close to the critical value of 1, dynamics is neutral, and X hovers around 0. As O is increased, dynamics becomes chaotic (X > 0); as O is decreased, dynamics becomes attractive (X < 0). B, The distribution of transmission probabilities emanating from each unit also influences dynamics. Three different types of units are shown, representing the three different types of exponential distributions that were examined. Thick arrows represent high transmission probabilities. The top unit shows transmission probabilities when the exponent B is low and the distribution is homogeneous. In this case, each unit acts to disperse trajectories, causing chaotic dynamics. The middle unit corresponds to intermediate values of B, where one to two transmission probabilities are strong. Here, each unit acts to focus trajectories, but with some dispersion, causing neutral dynamics. The lower unit illustrates the highly skewed distribution caused by large values of B. Here one connection dominates and all the rest are essentially zero. Units with high B distributions act to focus trajectories, leading to attractive dynamics. Figure 8 A is modified from Haldeman and Beggs, 2005, copyright American Physical Society, and is reproduced with permission by their approach, we here use an exponential distribution whose sharpness can be tuned through an exponent B, and we explore how B affects X. In these simulations, we use a network with 64 units that has 8 connections per unit. Qualitatively similar results obtain for networks with 64 connections per unit, suggesting that these findings are quite general. For small B (0 < B < 1.0), distributions are nearly flat and each connection has roughly the same probability of transmission. In this case, activity coming in to a unit will be spread widely and randomly to other connected units. This tends to disperse trajectories and leads to chaotic dynamics where X > 0. For intermediate values of B(1.2 < B < 1.8), one or two connections have transmission probabilities that are much larger than all the rest. Here activity coming in to a unit will tend to be transmitted to only one or two other units. This leads to propagation in which there is a balance of spreading and focus. While there is some variability in the paths that trajectories take, there is one path that is traveled most of the time. On average, dynamics tend to be neutral and X « 0. For large values of B(1.8 < B), one of the transmission probabilities is very near 1, while all of the others are near zero. So while a unit may receive convergent activation from two other units in the previous time step, it will almost always activate only one unit in the next time step. Under these conditions, units serve to bring different trajectories together, thus reducing distances over time and causing attractive dynamics with X < 0. Together, these simulations show that the distribution of connection strengths can also set the dynamics of a network (Fig. 8B).

How do changes in the number of connections affect dynamics? Although not directly in the field of neural networks, Stuart Kauffman has pursued this question in network models of gene regulatory networks. Since his studies are very likely to be relevant to our topic, we briefly mention them here. Kauff-man and colleagues (Kauffman S, 1969; Kauffman S et al., 2003; Kauffman SA and S Johnsen, 1991) examine networks where each binary unit can be either on (1) or off (0), and where each unit performs some Boolean function (e.g., AND, OR) on its inputs. Units are connected randomly, and the number of connections into each unit is determined by an order parameter K. Kauffman shows in these random Boolean networks that when K > 3, trajectories are very sensitive to small perturbations and dynamics is chaotic. When K = 2, however, trajectories are stable with respect to perturbations and the networks appear to operate at a critical point (Bornholdt S and T Rohlf, 2000). For K < 2, nearly all trajectories quickly fall into attractors. Kauffman and others (Gutowitz H and C Langton, 1995) have suggested that K governs a phase transition in these networks as it controls their dynamics. In some ways, high K networks may be similar to the neural network model described above when the distribution exponent B is small and all transmission probabilities are nearly equal. For intermediate values of B, one or two transmission probabilities are strong, and this may correspond to the critical case where K = 2 in Kauffman's networks. These possible connections are intriguing and deserve further exploration.

But why should dynamics matter? The dynamical regime of a network can strongly influence the types of computations it is able to perform (Vogels TP et al., 2005). Many models and experiments suggest that local networks support attractive dynamics (Amit Y and M Mascaro, 2001; Brunel N, 2000; Hopfield JJ, 1982; Jin DZ, 2002; Seung HS, 1998; Wills TJ et al., 2005). As mentioned earlier, strongly attractive dynamics is naturally good for setting up attractor states in which long-term memories can be stably stored. Such dynamics is also desirable for pattern completion, since a fragment of a stored pattern can be used as a cue to get the network into a state where it is near a basin of attraction and likely to evolve into the stored memory configuration. Moreover, attractive dynamics supports computations that favor categorization since they cause different stimuli to be grouped into the same response. For example, if a Wolfhound, a Chihuahua and a Beagle were all represented by positions in state space, attractive dynamics could cause trajectories from these points to all flow together, making it easy to set up the category of "dog." But the stability conferred by attractive dynamics also makes it difficult to steer trajectories away from strong attractors. Networks dominated by attractive dynamics would seem to lack flexibility.

In contrast, chaotic dynamics supports computations that favor discrimination since subtle differences in stimuli can produce widely different responses. Here too, there are a number of models and experiments that suggest that chaotic dynamics are prevalent in the brain (Aitken PG et al., 1995; Babloyantz A and A Destexhe, 1986; Breakspear M et al., 2003; Freeman WJ, 1994; Schiff SJ et al., 1994; van Vreeswijk C and H Sompolinsky, 1996). This dynamics could be useful in sensory systems where there is a great need to notice details of the incoming information stream. For example, whether a rabbit stays and eats or rapidly flees may be determined by only a few blades of grass in the visual field that seem to be moving in an unusual way. There have also been proposals that chaotic processing units could be used to perform logical or arithmetic computations since such units are naturally nonlinear (Sinha S and WL Ditto, 1999). However, networks with trajectories that rapidly diverge are unstable unless they are controlled.

With neutral dynamics, differences in inputs produce commensurate differences in responses. Not surprisingly, there are models and experiments that suggest this type of dynamics is used too (Beggs JM and D Plenz, 2003; Bertschinger N and T Natschlager, 2004; Haldeman C and JM Beggs, 2005; Latham PE and S Nirenberg, 2004; Maass W et al., 2002). This dynamics supports computations that favor efficient information transmission since a one-to-one mapping between stimuli and responses is maintained. They may also be optimal for information storage (Beggs JM and D Plenz, 2004; Haldeman C and JM Beggs, 2005). Several researchers have pointed out that neutral dynamics, "at the edge of chaos," may also be best for performing the widest variety of computations because it combines some of the variety of chaos with some of the stability of attractive systems (Bertschinger N and T Natschlager, 2004; Beggs 2007). It is argued that useful computations require both nonlinear transformations and stable representations of information. Perhaps neocortex, which is essential for higher-level computations, has largely neutral dynamics (Maass W et al., 2002; Natschlager T and W Maass, 2005).

To advance research in this area it will be necessary to form a tighter link between models and experiments. Many of the ideas about how connectivity influences dynamics described above have not yet been tested in living neural networks. Since nature often defies our expectations, it is essential that we develop better ways of interrogating networks of neurons. With advances in technology in the next ten years (Frechette ES et al., 2005), it may be possible to stimulate and record from thousands of neurons for periods of weeks at a time. The huge data sets that are likely to be produced will hopefully allow us to map the state space of living neural networks more closely.

It will also be important to investigate how different network topologies (e.g., random, small-world, scale-free) explicitly influence dynamics. The simulations described above treated all nodes in the network equiva-lently, but this is certainly a simplification. What happens when some nodes have different branching parameters and transmission probabilities than others? What if some nodes have more connections than others? These issues are only now beginning to be explored (Fox JJ and CC Hill, 2001), as the network topology of the brain at the local network level (Netoff TI et al., 2004; Song S et al., 2005) and at the large scale level (Achard S et al., 2006; Eguiluz VM et al., 2005; Sporns O et al., 2005; Sporns O and JD Zwi, 2004; Stam CJ et al., 2006) is still not well known. The connectivity patterns, and therefore the dynamics, at these different levels may not necessarily be the same (Breakspear M and CJ Stam, 2005; Jirsa VK, 2004).

Another area that deserves much attention is the relationship between dynamics and connectivity: How does brain activity, both acutely and chronically, alter the connectivity of neural networks? While activity-dependent synaptic plasticity has been extensively studied, most of this work has centered on how stimulation at one or a few synapses influences synaptic efficacy There is a need to expand the focus to explore how activity at the local network level may influence synaptic plasticity. In vivo, transmission at a single synapse is embedded in the context of rich background activity that is very influential (Leger JF et al., 2005). From this perspective, functional connectivity is very dynamic and may be different from the underlying structural connectivity (Sporns O et al., 2000). Since it has been shown that large-scale network connectivity can change from wakefulness to sleep (Massimini M et al., 2005), it seems likely that it would also change during transitions to other brain states as well, like seizures. Similar changes at the local network level should also be investigated. While it may be difficult to disentangle the contributions of connectivity and dynamics in these situations, their complexity suggests that these situations will be interesting and fruitful areas for further research.

In the previous sections we have shown how early models of memory storage in local recurrent networks led many to search for attractors in neurophys-iological data. While numerous examples of reproducible activity patterns in living neural networks have been found, very few experimental studies have addressed the dynamics of these networks quantitatively. By measuring the Lyapunov exponent in simple network models, it has become clear that network connectivity can profoundly influence dynamics. Experimental work in the future will hopefully begin to quantitatively address the dynamics of local cortical networks, perhaps even revealing how trajectories in cortical columns perform computations that form the building blocks of cognition.

Connectivity and Dynamics in Local Cortical Networks 111

Acknowledgements

This work was supported by the National Science Foundation and Indiana University.

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