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Random generalized Lotka–Volterra model

From Wikipedia, the free encyclopedia
Model in theoretical ecology and statistical mechanics
Example dynamics in the unique fixed point and multiple attractors phases withS=64{\displaystyle S=64} species. For both simulations(μα,γ,K,σK)=(4,0,1,0){\displaystyle (\mu _{\alpha },\gamma ,K,\sigma _{K})=(4,0,1,0)}. For UFP,σα=1{\displaystyle \sigma _{\alpha }=1}; for MA,σα=2{\displaystyle \sigma _{\alpha }=2}.

Therandom generalized Lotka–Volterra model (rGLV) is anecological model andrandom set of coupled ordinary differential equations where the parameters of thegeneralized Lotka–Volterra equation are sampled from aprobability distribution, analogously toquenched disorder. The rGLV models dynamics of a community of species in which each species' abundance grows towards acarrying capacity but is depleted due tocompetition from the presence of other species. It is often analyzed in themany-species limit using tools fromstatistical physics, in particular fromspin glass theory.

The rGLV has been used as a tool to analyzeemergent macroscopic behavior inmicrobial communities with dense, strong interspecies interactions. The model has served as a context for theoretical investigations studyingdiversity-stability relations in community ecology[1] and properties of static and dynamiccoexistence.[2][3] Dynamical behavior in the rGLV has been mapped experimentally in community microcosms.[4] The rGLV model has also served as an object of interest for the spin glass and disordered systems physics community to develop new techniques and numerical methods.[5][6][7][8][9]

Definition

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The random generalized Lotka–Volterra model is written as the system ofcoupled ordinary differential equations,[1][2][4][10]dNidt=riKiNi(KiNij(i)αijNj),i=1,,S,{\displaystyle {\frac {\mathrm {d} N_{i}}{\mathrm {d} t}}={\frac {r_{i}}{K_{i}}}N_{i}\left(K_{i}-N_{i}-\sum _{j(\neq i)}\alpha _{ij}N_{j}\right),\qquad i=1,\dots ,S,}whereNi{\displaystyle N_{i}} is the abundance of speciesi{\displaystyle i},S{\displaystyle S} is the number of species,Ki{\displaystyle K_{i}} is the carrying capacity of speciesi{\displaystyle i} in the absence of interactions,ri{\displaystyle r_{i}} sets a timescale, andα{\displaystyle \alpha } is arandom matrix whose entries arerandom variables with meanαij=μα/S{\displaystyle \langle \alpha _{ij}\rangle =\mu _{\alpha }/S}, variancevar(αij)=σα2/S{\displaystyle \mathrm {var} (\alpha _{ij})=\sigma _{\alpha }^{2}/S}, andcorrelationscorr(αij,αji)=γ{\displaystyle \mathrm {corr} (\alpha _{ij},\alpha _{ji})=\gamma } forij{\displaystyle i\neq j} where1γ1{\displaystyle -1\leq \gamma \leq 1}. Theinteraction matrix,α{\displaystyle \alpha }, may be parameterized as,αij=μαS+σαSaij,{\displaystyle \alpha _{ij}={\frac {\mu _{\alpha }}{S}}+{\frac {\sigma _{\alpha }}{\sqrt {S}}}a_{ij},}whereaij{\displaystyle a_{ij}} are standard random variables (i.e., zero mean and unit variance) withaijaji=γ{\displaystyle \langle a_{ij}a_{ji}\rangle =\gamma } forij{\displaystyle i\neq j}. The matrix entries may have any distribution with common finite first and second moments and will yield identical results in the largeS{\displaystyle S} limit due to thecentral limit theorem. The carrying capacities may also be treated as random variables withKi=K,var(Ki)=σK2.{\displaystyle \langle K_{i}\rangle =K,\,\operatorname {var} (K_{i})=\sigma _{K}^{2}.} Analyses by statistical physics-inspired methods have revealedphase transitions between different qualitative behaviors of the model in themany-species limit. In some cases, this may include transitions between the existence of aunique globally-attractive fixed point andchaotic, persistent fluctuations.

Steady-state abundances in the thermodynamic limit

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In thethermodynamic limit (i.e., the community has a very large number of species) where a unique globally-attractive fixed point exists, the distribution of species abundances can be computed using thecavity method while assuming the system isself-averaging. The self-averaging assumption means that the distribution of any one species' abundance between samplings of model parameters matches the distribution of species abundances within a single sampling of model parameters. In the cavity method, an additionalmean-field speciesi=0{\displaystyle i=0} is introduced and the response of the system is approximated linearly. The cavity calculation yields aself-consistent equation describing the distribution of species abundances as a mean-field random variable,N0{\displaystyle N_{0}}. WhenσK=0{\displaystyle \sigma _{K}=0}, the mean-field equation is,[1]0=N0(KμαmN0+q(μα2+γσα2)Z+σα2γχN0),{\displaystyle 0=N_{0}\left(K-\mu _{\alpha }m-N_{0}+{\sqrt {q\left(\mu _{\alpha }^{2}+\gamma \sigma _{\alpha }^{2}\right)}}Z+\sigma _{\alpha }^{2}\gamma \chi N_{0}\right),}wherem=N0,q=N02,χ=N0/K0{\displaystyle m=\langle N_{0}\rangle ,\,q=\langle N_{0}^{2}\rangle ,\,\chi =\langle \partial N_{0}/\partial K_{0}\rangle }, andZN(0,1){\displaystyle Z\sim {\mathcal {N}}(0,1)} is astandard normal random variable. Onlyecologically uninvadable solutions are taken (i.e., the largest solution forN0{\displaystyle N_{0}} in the quadratic equation is selected). The relevant susceptibility and moments ofN0{\displaystyle N_{0}}, which has atruncated normal distribution, are determined self-consistently.

Dynamical phases

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In the thermodynamic limit where there is an asymptotically large number of species (i.e.,S{\displaystyle S\to \infty }), there are three distinctphases: one in which there is a unique fixed point (UFP), another with a multipleattractors (MA), and a third with unbounded growth. In the MA phase, depending on whether species abundances are replenished at a small rate, may approach arbitrarily small population sizes, or are removed from the community when the population falls below some cutoff, the resulting dynamics may be chaotic with persistent fluctuations or approach an initial conditions-dependent steady state.[1]

The transition from the UFP to MA phase is signaled by the cavity solution becoming unstable to disordered perturbations. WhenσK=0{\displaystyle \sigma _{K}=0}, thephase transition boundary occurs when the parameters satisfy,σα=21+γ.{\displaystyle \sigma _{\alpha }={\frac {\sqrt {2}}{1+\gamma }}.}In theσK>0{\displaystyle \sigma _{K}>0} case, the phase boundary can still be calculated analytically, but no closed-form solution has been found; numerical methods are necessary to solve the self-consistent equations determining the phase boundary.


The transition to the unbounded growth phase is signaled by the divergence ofN0{\displaystyle \langle N_{0}\rangle } as computed in the cavity calculation.

Dynamical mean-field theory

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The cavity method can also be used to derive adynamical mean-field theory model for the dynamics. The cavity calculation yields a self-consistent equation describing the dynamics as aGaussian process defined by the self-consistent equation (forσK=0{\displaystyle \sigma _{K}=0}),[8]dN0dt=N0(t)[K0N0(t)μαm(t)σαη(t)+γσα20tdtχ(t,t)N0(t)],{\displaystyle {\frac {\mathrm {d} N_{0}}{\mathrm {d} t}}=N_{0}(t)\left[K_{0}-N_{0}(t)-\mu _{\alpha }m(t)-\sigma _{\alpha }\eta (t)+\gamma \sigma _{\alpha }^{2}\int _{0}^{t}\mathrm {d} t'\,\chi (t,t')N_{0}(t')\right],}wherem(t)=N0(t){\displaystyle m(t)=\langle N_{0}(t)\rangle },η{\displaystyle \eta } is a zero-mean Gaussian process withautocorrelationη(t)η(t)=N0(t)N0(t){\displaystyle \langle \eta (t)\eta (t')\rangle =\langle N_{0}(t)N_{0}(t')\rangle }, andχ(t,t)=δN0(t)/δK0(t)|K0(t)=K0{\displaystyle \chi (t,t')=\langle \left.\delta N_{0}(t)/\delta K_{0}(t')\right|_{K_{0}(t')=K_{0}}\rangle } is the dynamical susceptibility defined in terms of afunctional derivative of the dynamics with respect to a time-dependent perturbation of the carrying capacity.

Using dynamical mean-field theory, it has been shown that at long times, the dynamics exhibit aging in which the characteristic time scale defining the decay of correlations increases linearly in the duration of the dynamics. That is,CN(t,t+tτ)f(τ){\displaystyle C_{N}(t,t+t\tau )\to f(\tau )} whent{\displaystyle t} is large, whereCN(t,t)=N(t)N(t){\displaystyle C_{N}(t,t')=\langle N(t)N(t')\rangle } is the autocorrelation function of the dynamics andf(τ){\displaystyle f(\tau )} is a common scaling collapse function.[8][11]

When a small immigration rateλ1{\displaystyle \lambda \ll 1} is added (i.e., a small constant is added to the right-hand side of the equations of motion) the dynamics reach atime transitionally invariant state. In this case, the dynamics exhibit jumps betweenO(1){\displaystyle O(1)} andO(λ){\displaystyle O(\lambda )} abundances.[12]

Related articles

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References

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  1. ^abcdBunin, Guy (2017-04-28)."Ecological communities with Lotka-Volterra dynamics".Physical Review E.95 (4) 042414.Bibcode:2017PhRvE..95d2414B.doi:10.1103/PhysRevE.95.042414.PMID 28505745.
  2. ^abServán, Carlos A.; Capitán, José A.; Grilli, Jacopo; Morrison, Kent E.; Allesina, Stefano (August 2018)."Coexistence of many species in random ecosystems".Nature Ecology & Evolution.2 (8):1237–1242.Bibcode:2018NatEE...2.1237S.doi:10.1038/s41559-018-0603-6.ISSN 2397-334X.PMID 29988167.S2CID 49668570.
  3. ^Pearce, Michael T.; Agarwala, Atish; Fisher, Daniel S. (2020-06-23)."Stabilization of extensive fine-scale diversity by ecologically driven spatiotemporal chaos".Proceedings of the National Academy of Sciences.117 (25):14572–14583.Bibcode:2020PNAS..11714572P.doi:10.1073/pnas.1915313117.ISSN 0027-8424.PMC 7322069.PMID 32518107.
  4. ^abHu, Jiliang; Amor, Daniel R.; Barbier, Matthieu; Bunin, Guy; Gore, Jeff (2022-10-07)."Emergent phases of ecological diversity and dynamics mapped in microcosms".Science.378 (6615):85–89.Bibcode:2022Sci...378...85H.doi:10.1126/science.abm7841.ISSN 0036-8075.PMID 36201585.S2CID 240251815.
  5. ^Sidhom, Laura; Galla, Tobias (2020-03-02)."Ecological communities from random generalized Lotka-Volterra dynamics with nonlinear feedback".Physical Review E.101 (3) 032101.arXiv:1909.05802.Bibcode:2020PhRvE.101c2101S.doi:10.1103/PhysRevE.101.032101.hdl:10261/218552.PMID 32289927.S2CID 214667872.
  6. ^Biroli, Giulio; Bunin, Guy; Cammarota, Chiara (August 2018). "Marginally stable equilibria in critical ecosystems".New Journal of Physics.20 (8): 083051.arXiv:1710.03606.Bibcode:2018NJPh...20h3051B.doi:10.1088/1367-2630/aada58.ISSN 1367-2630.
  7. ^Ros, Valentina; Roy, Felix; Biroli, Giulio; Bunin, Guy; Turner, Ari M. (2023-06-21)."Generalized Lotka-Volterra Equations with Random, Nonreciprocal Interactions: The Typical Number of Equilibria".Physical Review Letters.130 (25) 257401.arXiv:2212.01837.Bibcode:2023PhRvL.130y7401R.doi:10.1103/PhysRevLett.130.257401.PMID 37418712.S2CID 254246297.
  8. ^abcRoy, F; Biroli, G; Bunin, G; Cammarota, C (2019-11-29)."Numerical implementation of dynamical mean field theory for disordered systems: application to the Lotka–Volterra model of ecosystems".Journal of Physics A: Mathematical and Theoretical.52 (48): 484001.arXiv:1901.10036.Bibcode:2019JPhA...52V4001R.doi:10.1088/1751-8121/ab1f32.ISSN 1751-8113.S2CID 59336358.
  9. ^Arnoulx de Pirey, Thibaut; Bunin, Guy (2024-03-05)."Many-Species Ecological Fluctuations as a Jump Process from the Brink of Extinction".Physical Review X.14 (1) 011037.arXiv:2306.13634.doi:10.1103/PhysRevX.14.011037.
  10. ^Altieri, Ada; Roy, Felix; Cammarota, Chiara; Biroli, Giulio (2021-06-23)."Properties of Equilibria and Glassy Phases of the Random Lotka-Volterra Model with Demographic Noise".Physical Review Letters.126 (25) 258301.arXiv:2009.10565.Bibcode:2021PhRvL.126y8301A.doi:10.1103/PhysRevLett.126.258301.hdl:11573/1623024.PMID 34241496.S2CID 221836142.
  11. ^Arnoulx de Pirey, Thibaut; Bunin, Guy (2024-03-05)."Many-Species Ecological Fluctuations as a Jump Process from the Brink of Extinction".Physical Review X.14 (1) 011037.arXiv:2306.13634.doi:10.1103/PhysRevX.14.011037.
  12. ^Arnoulx de Pirey, Thibaut; Bunin, Guy (2024-03-05)."Many-Species Ecological Fluctuations as a Jump Process from the Brink of Extinction".Physical Review X.14 (1) 011037.arXiv:2306.13634.doi:10.1103/PhysRevX.14.011037.

Further reading

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