Summary: What We Have Learned So Far

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Summary: What We Have Learned So Far small-world phenomenon Real-world networks: { Short path lengths High clustering Broad degree distributions, often power laws P (k) k γ Erdös-Renyi model: Short path lengths Poisson degrees No clustering Watts-Strogatz Small World model: short path lengths high clustering almost constant degrees Barabási-Albert scale-free model: short path lengths power-law degrees no clustering, but simple variants fix this

The Barabási-Albert Scale Free Model A model of network growth Based on the principle of preferential attachment - rich get richer! Yields networks with a powerlaw degree distribution 1. Take a small seed network, e.g. a few connected nodes 2. Let a new node of degree m enter the network 3. Connect the new node to existing nodes such that the probability of connecting to node i of degree ki is (average degree <k>=2m) 4. Repeat 2.-3. until N nodes.

Scale-Free Network Models: Summary For growing networks, preferential attachment yields power-law degree distributions The fundamental model: Barabási- Albert, where To be exact, it has to be linear: (If superlinear, winner takes it all and in the end one node has ALL the links! If sublinear, we get a stretched exponential degree distribution) Several mechanisms lead to the preferential attachment principle! (if mixed, e.g. combination of linear preferential and random attachment, we get exponents larger than 3!)

More Network Characteristics & Network Analysis

Degree Correlations If the degree of a vertex is k, does this affect the degrees of its neighbours? We could investigate the conditional probability of the neighbour having degree k In practice this is cumbersome to calculate (esp. in data analysis) Hence the average nearestneighbour degree knn(k) is typically used k nn (k) = 1 1 N k k i i,k i =k j ν i k j

Degree Correlations In practical network analysis, knn(k) is simply calculated by averaging over all neighbour degrees for each k Alternative method: the assortativity coefficient (Pearson correlation coefficient) Assortativity: positive degree correlations, knn(k) increasing with k Disassortativity: negative degree correlations, knn(k) decreasing with k r positive: assortative mixing r negative: disassortative mixing

Degree Correlations: Visualized Example maximal correlations no correlations maximal anticorrelations Trusina et al, PRL 92, 2004 Three networks with same degree sequence, differently rewired

Degree Correlations: Examples Social network based on mobile telephone calls S. Cerevisiae protein interactions and genetic regulation avg neighbour degree Onnela et al Maslov, Sneppen, Science 296 (2002) degree k interactions regulatory network Social networks are almost always assortative, biological networks disassortative

knn(k) for uncorrelated networks Let a first calculate the probability that a vertex of degree k is connected to a vertex of degree k Now In uncorrelated networks, this is equal to the probability that the vertex in the other end of a random link has degree k : there there are are P(k )N P(k )N vertices vertices of of degree degree k, k, and and k k times times this this number number of of edge edge ends ends points points to to them them there there are are altogether altogether N<k> N<k> edge edge ends ends

knn(k) for the BA model 1. 1. Take Take a small small seed seed network network -- eg eg 4 connected connected vertices vertices 2. 2. Create Create a new new vertex vertex with with m edges edges First we need an expression for the degree of vertex i at time t Let s write the rate equation: 3. 3. Connect Connect the the m edges edges to to existing existing vertices vertices with with a probability probability proportional proportional to to their their degree degree k, k, ie ie the the probability probability ππ i of i of choosing choosing vertex vertex i i is is 4. 4. Repeat Repeat 2.-3. 2.-3. until until the the network network has has grown grown to to desired desired size size of of N vertices vertices

knn(k) for the BA model Now let s solve this by integration Since (i = time of birth of vertex i) Let us now assume that as far as correlations go, BA networks correspond to random nets: (<k>=2m) we finally get

More correlation measures: the rich-club coefficient How connected are high-degree vertices among themselves? The rich-club coefficient measures how many edges E>k exist among the N>k vertices of degrees higher than k, divided by the maximum possible number N>k(N>k-1)/2 It is useful to compare the values against randomized reference networks Randomized reference: rewire the network whilst retaining its degree sequence, getting rid of correlations (the configuration model)

Rich-club coefficient: examples Colizza et al., Nature Physics 2, 2006 ratio of φ(k) in orig. nets to φ(k) in randomized counterparts

Betweenness Centrality Measures the amount of flow through a vertex (or an edge), if each vertex sends e.g. a signal through all other vertices via shortest paths Formally: number of shortest paths going through vertex/edge, such that the contribution of each path is divided by its multiplicity (if any) Computationally demanding, for a good algorithm see M. E. J. Newman, Phys. Rev. E 64, 016132 (2001) +1/2 +1 +1 +1/2 +1

Betweenness Centrality: An Example highest betweenness centrality Marriages between influential families in 13th century Florence

Betweenness Centrality: An Example Sample from a large social network based on telephone calls Betweenness calculated for edges: red=high

Closeness & Eigenvector Centrality Closeness: measures how far, on the average, a vertex is from all other vertices: where d ij is the distance along links from i to j Eigenvector: assigns relative scores to all nodes in the network based on the principle that connections to nodes having a high score contribute more to the score of the node in question A measure of influence

PageRank P R(i) = 1 d N + d j ν i P R(j) k out (j) PR(i) = PageRank of i, d = damping factor, N = number of pages, νi= in-neighbourhood of i In essence, a damped version of eigenvector centrality Corresponds to a random surfer following hyperlinks, who continues with probability d (assumed for Google to be 0.85), and jumps to a random page with probability 1-d

PageRank The Anatomy of a Large-Scale Hypertextual Web Search Engine Brin, S. and Page, L. (1998) The Anatomy of a Large-Scale Hypertextual Web Search Engine. In: Seventh International World-Wide Web Conference (WWW 1998), April 14-18, 1998, Brisbane, Australia.

Subgraphs & Motifs Subgraph: any set of nodes in the network, and the edges connecting them directed subgraphs of order 3 Clique: a fully connected subgraph k-clique: clique with k vertices cliques Motif: subgraph occurring in a network at a number significantly higher than in randomized counterpart k=3 k=4 k=5 k=6 randomization: pick two random links exchange endpoints repeat

Counting Motifs: Z-score number of times the subgraph M occurs in an empirical network average number of times the subgraph M in randomized reference ensemble standard deviation of n M in reference ensemble

Motifs: Example Milo et al., Science 303, 1538 (2004)

Motifs: Conservation in Evolution Yeast (S. Cerevisiae) protein interactions Find out motifs in the interaction network Find out orthologs for proteins in higher eucaryotes (eg humans) Calculate fraction of motifs where each protein has an ortholog Wuchty et al., Nature Genetics 35, 176 (2003)

Weighted Networks

Weighted networks Elements vertices Interactions edges An edge between v i and v j means v i and v j interact In reality, interactions can have different strengths, leading to weighted networks Vertex Edge Weight person friendship closeness neuron synapse synaptic strength WWW hyperlink none company ownership % owned gene regulation level of regulation