Models of DNA evolution

Models of DNA evolution

A number of different Markov models of DNA sequence evolution have been proposed. These substitution models differ in terms of the parameters used to describe the rates at which one nucleotide replaces another during evolution. These models are frequently used in molecular phylogenetic analyses. In particular, they are used during the calculation of likelihood of a tree (in Bayesian and maximum likelihood approaches to tree estimation) and they are used to estimate the evolutionary distance between sequences from the observed differences between the sequences. == Introduction == These models are phenomenological descriptions of the evolution of DNA as a string of four discrete states. These Markov models do not explicitly depict the mechanism of mutation nor the action of natural selection. Rather they describe the relative rates of different changes. For example, mutational biases and purifying selection favoring conservative changes are probably both responsible for the relatively high rate of transitions compared to transversions in evolving sequences. However, the Kimura (K80) model described below only attempts to capture the effect of both forces in a parameter that reflects the relative rate of transitions to transversions. Evolutionary analyses of sequences are conducted on a wide variety of time scales. Thus, it is convenient to express these models in terms of the instantaneous rates of change between different states (the Q matrices below). If we are given a starting (ancestral) state at one position, the model's Q matrix and a branch length expressing the expected number of changes to have occurred since the ancestor, then we can derive the probability of the descendant sequence having each of the four states. The mathematical details of this transformation from rate-matrix to probability matrix are described in the mathematics of substitution models section of the substitution model page. By expressing models in terms of the instantaneous rates of change we can avoid estimating a large numbers of parameters for each branch on a phylogenetic tree (or each comparison if the analysis involves many pairwise sequence comparisons). The models described on this page describe the evolution of a single site within a set of sequences. They are often used for analyzing the evolution of an entire locus by making the simplifying assumption that different sites evolve independently and are identically distributed. This assumption may be justifiable if the sites can be assumed to be evolving neutrally. If the primary effect of natural selection on the evolution of the sequences is to constrain some sites, then models of among-site rate-heterogeneity can be used. This approach allows one to estimate only one matrix of relative rates of substitution, and another set of parameters describing the variance in the total rate of substitution across sites. == DNA evolution as a continuous-time Markov chain == === Continuous-time Markov chains === Continuous-time Markov chains have the usual transition matrices which are, in addition, parameterized by time, t {\displaystyle t} . Specifically, if E 1 , E 2 , E 3 , E 4 {\displaystyle E_{1},E_{2},E_{3},E_{4}} are the states, then the transition matrix P ( t ) = ( P i j ( t ) ) {\displaystyle P(t)={\big (}P_{ij}(t){\big )}} where each individual entry, P i j ( t ) {\displaystyle P_{ij}(t)} refers to the probability that state E i {\displaystyle E_{i}} will change to state E j {\displaystyle E_{j}} in time t {\displaystyle t} . Example: We would like to model the substitution process in DNA sequences (i.e. Jukes–Cantor, Kimura, etc.) in a continuous-time fashion. The corresponding transition matrices will look like: P ( t ) = ( p A A ( t ) p A G ( t ) p A C ( t ) p A T ( t ) p G A ( t ) p G G ( t ) p G C ( t ) p G T ( t ) p C A ( t ) p C G ( t ) p C C ( t ) p C T ( t ) p T A ( t ) p T G ( t ) p T C ( t ) p T T ( t ) ) {\displaystyle P(t)={\begin{pmatrix}p_{\mathrm {AA} }(t)&p_{\mathrm {AG} }(t)&p_{\mathrm {AC} }(t)&p_{\mathrm {AT} }(t)\\p_{\mathrm {GA} }(t)&p_{\mathrm {GG} }(t)&p_{\mathrm {GC} }(t)&p_{\mathrm {GT} }(t)\\p_{\mathrm {CA} }(t)&p_{\mathrm {CG} }(t)&p_{\mathrm {CC} }(t)&p_{\mathrm {CT} }(t)\\p_{\mathrm {TA} }(t)&p_{\mathrm {TG} }(t)&p_{\mathrm {TC} }(t)&p_{\mathrm {TT} }(t)\end{pmatrix}}} where the top-left and bottom-right 2 × 2 blocks correspond to transition probabilities and the top-right and bottom-left 2 × 2 blocks corresponds to transversion probabilities. Assumption: If at some time t 0 {\displaystyle t_{0}} , the Markov chain is in state E i {\displaystyle E_{i}} , then the probability that at time t 0 + t {\displaystyle t_{0}+t} , it will be in state E j {\displaystyle E_{j}} depends only upon i {\displaystyle i} , j {\displaystyle j} and t {\displaystyle t} . This then allows us to write that probability as p i j ( t ) {\displaystyle p_{ij}(t)} . Theorem: Continuous-time transition matrices satisfy: P ( t + τ ) = P ( t ) P ( τ ) {\displaystyle P(t+\tau )=P(t)P(\tau )} Note: There is here a possible confusion between two meanings of the word transition. (i) In the context of Markov chains, transition is the general term for the change between two states. (ii) In the context of nucleotide changes in DNA sequences, transition is a specific term for the exchange between either the two purines (A ↔ G) or the two pyrimidines (C ↔ T) (for additional details, see the article about transitions in genetics). By contrast, an exchange between one purine and one pyrimidine is called a transversion. === Deriving the dynamics of substitution === Consider a DNA sequence of fixed length m evolving in time by base replacement. Assume that the processes followed by the m sites are Markovian independent, identically distributed and that the process is constant over time. For a particular site, let E = { A , G , C , T } {\displaystyle {\mathcal {E}}=\{A,\,G,\,C,\,T\}} be the set of possible states for the site, and p ( t ) = ( p A ( t ) , p G ( t ) , p C ( t ) , p T ( t ) ) {\displaystyle \mathbf {p} (t)=(p_{A}(t),\,p_{G}(t),\,p_{C}(t),\,p_{T}(t))} their respective probabilities at time t {\displaystyle t} . For two distinct x , y ∈ E {\displaystyle x,y\in {\mathcal {E}}} , let μ x y {\displaystyle \mu _{xy}\ } be the transition rate from state x {\displaystyle x} to state y {\displaystyle y} . Similarly, for any x {\displaystyle x} , let the total rate of change from x {\displaystyle x} be μ x = ∑ y ≠ x μ x y . {\displaystyle \mu _{x}=\sum _{y\neq x}\mu _{xy}\,.} The changes in the probability distribution p A ( t ) {\displaystyle p_{A}(t)} for small increments of time Δ t {\displaystyle \Delta t} are given by p A ( t + Δ t ) = p A ( t ) − p A ( t ) μ A Δ t + ∑ x ≠ A p x ( t ) μ x A Δ t . {\displaystyle p_{A}(t+\Delta t)=p_{A}(t)-p_{A}(t)\mu _{A}\Delta t+\sum _{x\neq A}p_{x}(t)\mu _{xA}\Delta t\,.} In other words, (in frequentist language), the frequency of A {\displaystyle A} 's at time t + Δ t {\displaystyle t+\Delta t} is equal to the frequency at time t {\displaystyle t} minus the frequency of the lost A {\displaystyle A} 's plus the frequency of the newly created A {\displaystyle A} 's. Similarly for the probabilities p G ( t ) {\displaystyle p_{G}(t)} , p C ( t ) {\displaystyle p_{C}(t)} and p T ( t ) {\displaystyle p_{T}(t)} . These equations can be written compactly as p ( t + Δ t ) = p ( t ) + p ( t ) Q Δ t , {\displaystyle \mathbf {p} (t+\Delta t)=\mathbf {p} (t)+\mathbf {p} (t)Q\Delta t\,,} where Q = ( − μ A μ A G μ A C μ A T μ G A − μ G μ G C μ G T μ C A μ C G − μ C μ C T μ T A μ T G μ T C − μ T ) {\displaystyle Q={\begin{pmatrix}-\mu _{A}&\mu _{AG}&\mu _{AC}&\mu _{AT}\\\mu _{GA}&-\mu _{G}&\mu _{GC}&\mu _{GT}\\\mu _{CA}&\mu _{CG}&-\mu _{C}&\mu _{CT}\\\mu _{TA}&\mu _{TG}&\mu _{TC}&-\mu _{T}\end{pmatrix}}} is known as the rate matrix. Note that, by definition, the sum of the entries in each row of Q {\displaystyle Q} is equal to zero. It follows that p ′ ( t ) = p ( t ) Q . {\displaystyle \mathbf {p} '(t)=\mathbf {p} (t)Q\,.} For a stationary process, where Q {\displaystyle Q} does not depend on time t, this differential equation can be solved. First, P ( t ) = exp ⁡ ( t Q ) , {\displaystyle P(t)=\exp(tQ),} where exp ⁡ ( t Q ) {\displaystyle \exp(tQ)} denotes the exponential of the matrix t Q {\displaystyle tQ} . As a result, p ( t ) = p ( 0 ) P ( t ) = p ( 0 ) exp ⁡ ( t Q ) . {\displaystyle \mathbf {p} (t)=\mathbf {p} (0)P(t)=\mathbf {p} (0)\exp(tQ)\,.} === Ergodicity === If the Markov chain is irreducible, i.e. if it is always possible to go from a state x {\displaystyle x} to a state y {\displaystyle y} (possibly in several steps), then it is also ergodic. As a result, it has a unique stationary distribution π = { π x , x ∈ E } {\displaystyle {\boldsymbol {\pi }}=\{\pi _{x},\,x\in {\mathcal {E}}\}} , where π x {\displaystyle \pi _{x}} corresponds to the proportion of time spent in state x {\displaystyle x} after the Markov chain has run for an infinite amount of time. In DNA evo

Radar geo-warping

Radar geo-warping is the adjustment of geo-referenced radar images and video data to be consistent with a geographical projection. This image warping avoids any restrictions when displaying it together with video from multiple radar sources or with other geographical data including scanned maps and satellite images which may be provided in a particular projection. There are many areas where geo warping has unique benefits: Single radar video signal displayed together with maps of different geographical projections. E.g. Mercator UTM stereographic Multiple radar video signals displayed simultaneously: Having the computing power to do so on one computer. Adapting the projection of all radar signals allowing the geographically correct display and accurate superimposition of those videos. Slant range correction: a modern 3D radar system can measure the height of a target and hence it is possible to correct the radar video by the real corrected range of the target. Slant Range Correction also allows to compensate the radar tower height e.g. for maritime surveillance radars. == Introduction == Radar video presents the echoes of electromagnetic waves a radar system has emitted and received as reflections afterwards. These echoes are typically presented on a computer screen with a color-coding scheme depicting the reflection strength. Two problems have to be solved during such a visualization process. The first problem arises from the fact that typically the radar antenna turns around its position and measures the reflection echo distances from its position in one direction. This effectively means that the radar video data are present in polar coordinates. In older systems the polar oriented picture has been displayed in so called plan position indicators (PPI). The PPI-scope uses a radial sweep pivoting about the center of the presentation. This results in a map-like picture of the area covered by the radar beam. A long-persistence screen is used so that the display remains visible until the sweep passes again. Bearing to the target is indicated by the target's angular position in relation to an imaginary line extending vertically from the sweep origin to the top of the scope. The top of the scope is either true north (when the indicator is operated in the true bearing mode) or ship's heading (when the indicator is operated in the relative bearing mode). For visualization on a modern computer screen the polar coordinates have to be converted into Cartesian coordinates. This process called radar scan conversion is presented with more detail in the next section. The second problem to solve arises from the fact that a radar system is placed in the real world and measures real world echo positions. These echoes have to be displayed together with other real world data like object positions, vector maps and satellite images in a consistent way. All this information refers to the curved earth surface but is displayed on a flat computer display. Building a link from real world earth positions to display pixels is commonly called geographical referencing or in short geo-referencing. Part of the geo-referencing process is to map the 3D earth surface onto a 2D display. This process of a geographical projection can be performed in many ways, but different data sources have their own 'natural' projection. E.g. Cartesian radar video data from a radar source on the earth surface are geo-referenced by a so-called radar projection. When using this radar projection the Cartesian radar video pixels can directly displayed on a computer screen (only being linearly transformed according to the current position on the screen and e.g. the current zoom level). A problem now arises if e.g. also a satellite map shall be shown together with the radar video data. The 'natural' geographical projection of a satellite image would be a satellite projection which depends on the satellite orbit, position and further parameters. Now either the satellite image has to be reprojected to a radar projection or the radar video has to use the satellite projection. This geographical re-projection is also called geographical warping or Geo Warping where each image pixel has to be transformed from one projection into another. This article describes in further detail the Geo Warping of radar video images in real time. It will also show that radar video Geo Warping is done most efficiently when it is integrated with the radar scan conversion process. == Radar-scan conversion == This section describes the principles of the radar-scan conversion (RSC) process. The radar supplies its measured data in polar coordinates (ρ,θ) directly from the rotating antenna. ρ defines the target/echo distance and θ the target angle in polar world coordinates. These data are measured, digitized and stored in a polar coordinate polar store or polar pixmap. The main RSC task is to convert these data to Cartesian (x, y) display coordinates, creating the necessary display pixels. The RSC process is influenced by the current zoom, shift and rotation settings defining which part of the 'world' shall be visible in the display image. As detailed later the RSC process also takes the currently used geographical projection into account when the radar video images are Geo Warped. The OpenGL RSC is implemented using a reverse scan conversion approach which calculates for every image pixel the most appropriate radar amplitude value in the polar store. This approach generates an optimal image without any artifacts known from forward spoke fill algorithms. By applying bi-linear filtering between adjacent pixels in the polar store during the conversion process the OpenGL RSC finally achieves a very high visual quality radar display image for every zoom level, creating smooth images of the radar echoes. == Radar projection == This section illustrates how radar video data are geo referenced and displayed on a computer screen. The radar sensor is positioned on the earth surface with a height h above the ground. It measures the direct distance d to the target (and not e.g. the distance the target is away from the radar if one would move on the earth surface). This distance is then used in the display plane after adjustment to the current display zoom level by the radar scan converter (RSC). Now it has to be clarified how the radar video data is geo referenced. This basically means, that if we want to display a geographical real world object (like e.g. a light house) which is at the same real world position as the radar target, that it also shall appear at the same position in the display plane. This is realized by calculating the distance from the radar sensor to the respective real world object and use that distance in the display plane. The position of the real world object is typically given in geographical coordinates (latitude, longitude and height above the earth surface). In other words, using a radar projection with geographical data is done by simulating a radar measurement process with the real world objects and use the resulting range and azimuth in the display plane. The second picture to the right shows an example radar projection with the center of projection (COP) at latitude 50.0° and longitude 0.0° which is also the radar position. The dashed lines are the equal-latitude and equal-longitude lines on top of the background map. The solid lines show equal-range and equal-azimuth with the respect to the radar position. It is a feature of the radar projection that equal-range lines are circles and equal-azimuth lines are straight lines. This is necessary to display radar video consistently with other map data when using a radar projection where the projection center has to be the radar position. == Geo Warping process == This section explains the actual geo warping or re-projection process when applied to radar video in real time. Assume we want to display radar video on top of a satellite image. As an example we use the CIB projection which is used to display satellite data in CIB (Controlled Image Base) format. The Figure Geo Warping Radar to CIB Projection shows dashed the maximal range circle for a range of 111 km or 60 miles using the radar projection. Such a range is typical for long range coastal surveillance radars. As stated in the last section this is a perfect circle also on the computer screen. The solid line ellipse shows the same range circle for the CIB projection. Typically the errors occurring without Geo Warping are smallest near the radar position if at least the projection center (COP) coincides with the radar position, as realized in our example. Otherwise the error distribution depends both on the used projection and also on the projection parameters. Thus, in our case the errors are most significant near the maximum radar range. The CIB projection error corrected in east–west direction at half the radar range is 2.6 km and is 5.3 km at the full radar range of 111 km. An error of 5.3 km is

Guideline execution engine

A guideline execution engine is a computer program which can interpret a clinical guideline represented in a computerized format and perform actions towards the user of an electronic medical record. A guideline execution engine needs to communicate with a host clinical information system. Virtual Medical Record (vMR) is one possible interface which can be used. The engine's main function is to manage instances of executed guidelines of individual patients. == Architecture == The following modules are generally needed for any engine: interface to clinical information system new guidelines loading module guideline interpreter module clinical events parser alert/recommendations dispatch == Guideline Interchange Format == The Guideline Interchange Format (GLIF) is a computer representation format for clinical guidelines. Represented guidelines can be executed using a guideline execution engine. The format has several versions as it has been improved. In 2003 GLIF3 was introduced. == Use of third party workflow engine as a guideline execution engine == Some commercial electronic health record systems use a workflow engine to execute clinical guidelines. RetroGuide and HealthFlow are examples of such an approach.

Parallel terraced scan

The parallel terraced scan is a multi-agent based search technique that is basic to cognitive architectures, such as Copycat, Letter-string, the Examiner, Tabletop, and others. It was developed by John Rehling and Douglas Hofstadter at the Center for Research on Concepts and Cognition at Indiana University, Bloomington. The parallel terraced scan builds on the concepts of the workspace, coderack, conceptual memory, and temperature. According to Hofstadter the parallel and random nature of the processing captures aspects of human cognition.

Tim Houlne

Tim Houlne is an American business executive, entrepreneur, and author known for his work in outsourcing and homeshoring, remote working, and artificial intelligence (AI) in customer service. He is the founder and CEO of Humach, a company that uses human agents and AI in customer experience solutions. Previously, he was co-founder and CEO of Working Solutions, a virtual contact center company in the United States. == Early life and education == Houlne graduated from Missouri Western State University (MWSU) in 1986 with a bachelor's degree in business administration and from the University of Texas in Dallas with an MBA. In 2024, MWSU and North Central Missouri College renamed the Convergent Technology Alliance Center to the Houlne Center for Convergent Technology. The 20,000 square-foot learning laboratory provides training and applied education experiences in industries such as AI, cybersecurity, manufacturing and construction, and service technologies. == Career == In 1998, Houlne co-founded Working Solutions, a Plano, Texas-based U.S. outsourcing company that provides customer service using remote, home-based agents. As CEO, he oversaw the development of a virtual workforce model that routes service calls to either domestic or offshore agents, according to client needs and service requirements. In 2015, Houlne founded Humach, a customer experience outsourcing provider that uses human service agents with AI-based digital agents. The company derives its name from the combination of services provided by humans and machines. Its clients include Amazon, Carfax and McDonald's. The company acquired InfiniteAI in 2020, and Markets EQ in 2025. In 2013, Houlne was named a finalist for the Ernst & Young Entrepreneur of the Year Award (Southwest Region).He is the co-author of several books focused on the evolution of work, the gig economy, and the influence of AI in customer-facing roles. == Works == The New World of Work: From the Cube to the Cloud (2013) ISBN 0982562276 OCLC 813933360 The New World of Work, Second Edition: The Cube, the Cloud and What's Next (2023) ISBN 9781642258318 OCLC 1389815847 The Intelligent Workforce: How Humans & Machines Will Co-Create a Better Future (2024) ISBN 9798887501604 OCLC 1439598569

Automatic scorer

An automatic scorer is the computerized scoring system to keep track of scoring in ten-pin bowling. It was introduced en masse in bowling alleys in the 1970s and combined with mechanical pinsetters to detect overturned pins. By eliminating the need for manual score-keeping, these systems have introduced new bowlers into the game who otherwise would not participate because they had to count the score themselves, as many do not understand the mathematical formula involved in bowler scoring. At first, people were skeptical about whether a computer could keep an accurate score. In the twenty-first century, automatic scorers are used in most bowling centers around the world. The three manufacturers of these specialty computers have been Brunswick Bowling, AMF Bowling (later QubicaAMF), and RCA. == History == Automatic equipment is considered a cornerstone of the modern bowling center. The traditional bowling center of the early 20th century was advanced in automation when the pinsetter person ("pin boy"), who set back up by hand the bowled down pins, was replaced by a machine that automatically replaced the pins in their proper play positions. This machine came out in the 1950s. A detection system was developed from the pinsetter mechanism in the 1960s that could tell which pins had been knocked down, and that information could be transferred to a digital computer. Automatic electronic scoring was first conceived by Robert Reynolds, who was described by a newspaper story at the time as "a West Coast electronics calculator expert." He worked with the technical staff of Brunswick Bowling to develop it. The goal was realized in the late 1960s when a specialized computer was designed for the purpose of automatic scorekeeping for bowling. The field test for the automatic scorer took place at Village Lanes bowling center, Chicago in 1967. The scoring machine received approval for official use by the American Bowling Congress in August of that year. They were first used in national official league gaming on October 10, 1967. In November, Brunswick announced that they were accepting orders for the new digital computer, which cost around $3,000 per bowling lane. Bowling centers that installed these new automatic scoring devices in the 1970s charged a ten cents extra per line of scoring for the convenience. == Description == Each Automatic Scorer computer unit kept score for four lanes. It had two bowler identification panels serving two lanes each. The bowler pushed it into his named position when his turn came up so the computer knew who was bowling and score accordingly. After the bowler rolled the bowling ball down the lane and knocked down pins, the pinsetter detected which pins were down and relayed this information back to the computer for scoring. The result was then printed on a scoresheet and projected overhead onto a large screen for all to see. The Automatic Scorer digital computer was mathematically accurate, however the detection system at the pinsetter mechanism sometimes reported the wrong number of pins knocked down. The computer could be corrected manually for any errors in the system; similarly, human errors, such as neglecting to move the bowler identification mechanism, could be corrected for by manual action. The scorer could take into account bowlers' handicaps and could adjust for late-arriving bowlers. The automatic scorer is directly connected to the foul detection unit. As a result, foul line violations are automatically scored. Brunswick had put ten years of research and development into the Automatic Scorer, and by 1972 there were over 500 of these computers installed in bowling centers around the world. AMF Bowling, competitor to Brunswick, entered into the automatic scorer computer field during the 1970s and their systems were installed into their brand of bowling centers. By 1974, RCA was also making these computers for automatic scoring. == Reception and further developments == The purposes of the computerized scoring were to avoid errors by human scorers and to prevent cheating. It had the side benefit of speeding up the progress of the game and introducing new bowlers to the game. Score-keeping for bowling is based on a formula that many new to bowling were not familiar with and thought difficult to learn. These casual bowlers unfamiliar with the formula thought the scores given by the computers were confusing. Some bowlers were not comfortable with automatic scorers when they were introduced in the 1970s, so kept score using the traditional method on paper score sheets. The introduction of this device increased the popularity of the sport. Automatic scorers came to be considered a normal part of modern bowling installations worldwide, with owners and managers saying that bowlers expect such equipment to be present in bowling establishments and that business increased following their introduction. Brunswick introduced a color television style automatic scorer in 1983. Bowling center owners could use these style automatic scorers for advertising, management, videos, and live television. By the 2010s, these types of electronic visual displays could show bowler avatars and social media connections to publicize the bowlers' scores. Some are capable of being extended entertainment systems of games for children and adults. Some scoring systems support variations on traditional bowling, such as different kinds of bingo games where certain pins have to be knocked down at certain times or practice regimes where certain spares have to be accomplished. By this point, QubicaAMF Worldwide, an outgrowth of AMF, was one of the leading providers of bowling scoring equipment.

Resilience (mathematics)

In mathematical modeling, resilience refers to the ability of a dynamical system to recover from perturbations and return to its original stable steady state. It is a measure of the stability and robustness of a system in the face of changes or disturbances. If a system is not resilient enough, it is more susceptible to perturbations and can more easily undergo a critical transition. A common analogy used to explain the concept of resilience of an equilibrium is one of a ball in a valley. A resilient steady state corresponds to a ball in a deep valley, so any push or perturbation will very quickly lead the ball to return to the resting point where it started. On the other hand, a less resilient steady state corresponds to a ball in a shallow valley, so the ball will take a much longer time to return to the equilibrium after a perturbation. The concept of resilience is particularly useful in systems that exhibit tipping points, whose study has a long history that can be traced back to catastrophe theory. While this theory was initially overhyped and fell out of favor, its mathematical foundation remains strong and is now recognized as relevant to many different systems. == History == In 1973, Canadian ecologist C. S. Holling proposed a definition of resilience in the context of ecological systems. According to Holling, resilience is "a measure of the persistence of systems and of their ability to absorb change and disturbance and still maintain the same relationships between populations or state variables". Holling distinguished two types of resilience: engineering resilience and ecological resilience. Engineering resilience refers to the ability of a system to return to its original state after a disturbance, such as a bridge that can be repaired after an earthquake. Ecological resilience, on the other hand, refers to the ability of a system to maintain its identity and function despite a disturbance, such as a forest that can regenerate after a wildfire while maintaining its biodiversity and ecosystem services. With time, the once well-defined and unambiguous concept of resilience has experienced a gradual erosion of its clarity, becoming more vague and closer to an umbrella term than a specific concrete measure. == Definition == Mathematically, resilience can be approximated by the inverse of the return time to an equilibrium given by resilience ≡ − Re ( λ 1 ( A ) ) {\displaystyle {\text{resilience}}\equiv -{\text{Re}}(\lambda _{1}({\textbf {A}}))} where λ 1 {\textstyle \lambda _{1}} is the maximum eigenvalue of matrix A {\textstyle {\textbf {A}}} . The largest this value is, the faster a system returns to the original stable steady state, or in other words, the faster the perturbations decay. == Applications and examples == In ecology, resilience might refer to the ability of the ecosystem to recover from disturbances such as fires, droughts, or the introduction of invasive species. A resilient ecosystem would be one that is able to adapt to these changes and continue functioning, while a less resilient ecosystem might experience irreversible damage or collapse. The exact definition of resilience has remained vague for practical matters, which has led to a slow and proper application of its insights for management of ecosystems. In epidemiology, resilience may refer to the ability of a healthy community to recover from the introduction of infected individuals. That is, a resilient system is more likely to remain at the disease-free equilibrium after the invasion of a new infection. Some stable systems exhibit critical slowing down where, as they approach a basic reproduction number of 1, their resilience decreases, hence taking a longer time to return to the disease-free steady state. Resilience is an important concept in the study of complex systems, where there are many interacting components that can affect each other in unpredictable ways. Mathematical models can be used to explore the resilience of such systems and to identify strategies for improving their resilience in the face of environmental or other changes. For example, when modelling networks it is often important to be able to quantify network resilience, or network robustness, to the loss of nodes. Scale-free networks are particularly resilient since most of their nodes have few links. This means that if some nodes are randomly removed, it is more likely that the nodes with fewer connections are taken out, thus preserving the key properties of the network.