- Open Access
Flow currents support simple and versatile trail-tracking strategies
Phys. Rev. Research 8, 013019 – Published 12 January, 2026
DOI: https://doi.org/10.1103/qj9q-p7rg
Abstract
Aquatic animals offer compelling evidence that flow sensing alone, without vision, is sufficient to guide a swimming organism to the source of an unsteady hydrodynamic trail. However, the sensory feedback strategies that allow these remarkable trail-tracking abilities remain opaque. Here, by integrating mechanistic flow simulations with reinforcement learning techniques, we discovered two simple and equally effective strategies for hydrodynamic trail following. Though not a priori obvious, these strategies possess parsimonious interpretations, analogous to Braitenberg’s simplest vehicles, where the agent senses local flow signals and turns away from or toward the direction of stronger signals. A rigorous stability analysis shows that the effectiveness of these strategies in robustly tracking flow currents is independent of the type of sensor but depends on sensor placement and the traveling nature of the flow signal. Importantly, these results inform a suite of versatile strategies for hydrodynamic trail following applicable to both vortical and turbulent flows. These insights support the future design and implementation of adaptive real-time sensory feedback strategies for autonomous robots in dynamic flow environments.
Physics Subject Headings (PhySH)
Article Text
Supplemental Material
References (132)
- E. M. Purcell, Life at low Reynolds number, in Physics and Our World: Reissue of the Proceedings of a Symposium in Honor of Victor F. Weisskopf (American Institute of Physics, Melville, New York, 1977), pp. 47–67.
- D. Roszak and R. Colwell, Survival strategies of bacteria in the natural environment, Microbiol. Rev. 51, 365 (1987).
- G. M. Viswanathan, M. G. Da Luz, E. P. Raposo, and H. E. Stanley, The Physics of Foraging: An Introduction to Random Searches and Biological Encounters (Cambridge University Press, Cambridge, 2011).
- National Academies of Sciences, Engineering, and Medicine, Physics of Life (The National Academies Press, Washington, DC, 2022), https://nap.nationalacademies.org/catalog/26403/physics-of-life.
- H. C. Berg, Random Walks in Biology (Princeton University Press, Princeton, New Jersey, 1993).
- H. C. Berg and E. M. Purcell, Physics of chemoreception, Biophys. J. 20, 193 (1977).
- J. Adler, Chemoreceptors in bacteria: Studies of chemotaxis reveal systems that detect attractants independently of their metabolism, Science 166, 1588 (1969).
- E. F. Keller and L. A. Segel, Model for chemotaxis, J. Theor. Biol. 30, 225 (1971).
- H. C. Berg and D. A. Brown, Chemotaxis in Escherichia coli analysed by three-dimensional tracking, Nature (London) 239, 500 (1972).
- A. Gomez-Marin, G. J. Stephens, and M. Louis, Active sampling and decision making in Drosophila chemotaxis, Nat. Commun. 2, 441 (2011).
- K. J. Leitch, F. V. Ponce, W. B. Dickson, F. van Breugel, and M. H. Dickinson, The long distance flight behavior of Drosophila supports an agent-based model for wind-assisted dispersal in insects, Proc. Natl. Acad. Sci. USA 118, e2013342118 (2021).
- N. Kadakia et al., Odour motion sensing enhances navigation of complex plumes, Nature (London) 611, 754 (2022).
- A. Mafra-Neto and R. T. Cardé, Fine-scale structure of pheromone plumes modulates upwind orientation of flying moths, Nature (London) 369, 142 (1994).
- R. T. Cardé and A. Mafra-Neto, Neuroethology of pheromone-mediated responses, Insect Pheromone Research, edited by R. T. Cardé and A. K. Minks (Springer, New York, 1997), pp. 275–290.
- M. J. Weissburg, M. Doall, and J. Yen, Following the invisible trail: Kinematic analysis of mate-tracking in the copepod Temora longicornis, Philos. Trans. R. Soc. London B 353, 701 (1998).
- J. Yen, J. K. Sehn, K. Catton, A. Kramer, and O. Sarnelle, Pheromone trail following in three dimensions by the freshwater copepod Hesperodiaptomus shoshone, J. Plankton Res. 33, 907 (2011).
- R. Monthiller, A. Loisy, M. A. Koehl, B. Favier, and C. Eloy, Surfing on turbulence: A strategy for planktonic navigation, Phys. Rev. Lett. 129, 064502 (2022).
- M. Vergassola, E. Villermaux, and B. I. Shraiman, ‘Infotaxis’ as a strategy for searching without gradients, Nature (London) 445, 406 (2007).
- S. H. Singh, F. van Breugel, R. P. Rao, and B. W. Brunton, Emergent behaviour and neural dynamics in artificial agents tracking odour plumes, Nat. Mach. Intell. 5, 58 (2023).
- K. Pohlmann, F. W. Grasso, and T. Breithaupt, Tracking wakes: The nocturnal predatory strategy of piscivorous catfish, Proc. Natl. Acad. Sci. USA 98, 7371 (2001).
- K. Pohlmann, J. Atema, and T. Breithaupt, The importance of the lateral line in nocturnal predation of piscivorous catfish, J. Exp. Biol. 207, 2971 (2004).
- J. M. Gardiner and J. Atema, Sharks need the lateral line to locate odor sources: Rheotaxis and eddy chemotaxis, J. Exp. Biol. 210, 1925 (2007).
- G. Dehnhardt, B. Mauck, and H. Bleckmann, Seal whiskers detect water movements, Nature (London) 394, 235 (1998).
- G. Dehnhardt, B. Mauck, W. Hanke, and H. Bleckmann, Hydrodynamic trail-following in harbor seals (Phoca vitulina), Science 293, 102 (2001).
- G. R. Spedding, Wake signature detection, Annu. Rev. Fluid Mech. 46, 273 (2014).
- B. Colvert, K. Chen, and E. Kanso, Local flow characterization using bioinspired sensory information, J. Fluid Mech. 818, 366 (2017).
- B. Colvert, G. Liu, H. Dong, and E. Kanso, Flowtaxis in the wakes of oscillating airfoils, Theor. Comput. Fluid Dyn. 34, 545 (2020).
- S. D. Boie et al., Information-theoretic analysis of realistic odor plumes: What cues are useful for determining location? PLoS Comput. Biol. 14, e1006275 (2018).
- J. Ježov, O. Akanyeti, L. D. Chambers, and M. Kruusmaa, Sensing oscillations in unsteady flow for better robotic swimming efficiency, in 2012 IEEE International Conference on Systems, Man, and Cybernetics (SMC) (Seoul, Korea, 2012), pp. 91–96.
- N. Muhammad et al., Map-based localization and loop-closure detection from a moving underwater platform using flow features, Auton. Robots 43, 1419 (2019).
- P. Gunnarson and J. O. Dabiri, Fish-inspired tracking of underwater turbulent plumes, Bioinspir. Biomim 19, 056024 (2024).
- P. Gunnarson and J. O. Dabiri, Surfing vortex rings for energy-efficient propulsion, Proc. Natl. Acad. Sci. USA 4, pgaf031 (2025).
- H. Bleckmann, J. Mogdans, and S. L. Coombs, Flow Sensing in Air and Water (Springer-Verlag, Berlin, Germany, 2014), Vol. 976.
- J. C. Liao, D. N. Beal, G. V. Lauder, and M. S. Triantafyllou, Fish exploiting vortices decrease muscle activity, Science 302, 1566 (2003).
- D. N. Beal, F. S. Hover, M. S. Triantafyllou, J. C. Liao, and G. V. Lauder, Passive propulsion in vortex wakes, J. Fluid Mech. 549, 385 (2006).
- L. Li et al., Vortex phase matching as a strategy for schooling in robots and in fish, Nat. Commun. 11, 5408 (2020).
- S. Dijkgraaf, The functioning and significance of the lateral-line organs, Biol. Rev. 38, 51 (1963).
- J. Blaxter, Structure and development of the lateral line, Biol. Rev. 62, 471 (1987).
- A. Kroese and N. Schellart, Velocity-and acceleration-sensitive units in the trunk lateral line of the trout, J. Neurophysiol. 68, 2212 (1992).
- J. Engelmann, W. Hanke, J. Mogdans, and H. Bleckmann, Hydrodynamic stimuli and the fish lateral line, Nature (London) 408, 51 (2000).
- S. Coombs, P. Görner, and H. Münz, The Mechanosensory Lateral Line: Neurobiology and Evolution (Springer Science and Business Media, New York, 2012).
- N. Schulte-Pelkum, S. Wieskotten, W. Hanke, G. Dehnhardt, and B. Mauck, Tracking of biogenic hydrodynamic trails in harbour seals (Phoca vitulina), J. Exp. Biol. 210, 781 (2007).
- S. P. Windsor, S. E. Norris, S. M. Cameron, G. D. Mallinson, and J. C. Montgomery, The flow fields involved in hydrodynamic imaging by blind Mexican cave fish (Astyanax fasciatus). Part I: Open water and heading towards a wall, J. Exp. Biol. 213, 3819 (2010).
- M. J. McHenry, J. A. Strother, and S. M. Van Netten, Mechanical filtering by the boundary layer and fluid–structure interaction in the superficial neuromast of the fish lateral line system, J. Comp. Physiol. A 194, 795 (2008).
- M. E. McConney et al., Biologically inspired design of hydrogel-capped hair sensors for enhanced underwater flow detection, Soft Matter 5, 292 (2009).
- H. Beem, M. Hildner, and M. Triantafyllou, Calibration and validation of a harbor seal whisker-inspired flow sensor, Smart Mater. Struct. 22, 014012 (2013).
- A. C. Hollenbeck, R. Grandhi, J. H. Hansen, and A. M. Pankonien, Bioinspired artificial hair sensors for flight-by-feel of unmanned aerial vehicles: A review, AIAA J. 61, 5206 (2023).
- A. D. Hasler and A. T. Scholz, Olfactory Imprinting and Homing in Salmon: Investigations into the Mechanism of the Imprinting Process (Springer Science and Business Media, New York, 2012).
- E. Zermelo, Über das Navigationsproblem bei ruhender oder veränderlicher Windverteilung, ZAMM—J. App. Math. Mech./Z. Angew. Math. Mech. 11, 114 (1931).
- P. Gunnarson, I. Mandralis, G. Novati, P. Koumoutsakos, and J. O. Dabiri, Learning efficient navigation in vortical flow fields, Nat. Commun. 12, 7143 (2021).
- Y. Jiao, H. Hang, J. Merel, and E. Kanso, Sensing flow gradients is necessary for learning autonomous underwater navigation, Nat. Commun. 16, 3044 (2025).
- K. Krishna, Z. Song, and S. L. Brunton, Finite-horizon, energy-efficient trajectories in unsteady flows, Proc. R. Soc. A 478, 20210255 (2022).
- L. Biferale, F. Bonaccorso, M. Buzzicotti, P. Clark Di Leoni, and K. Gustavsson, Zermelo’s problem: Optimal point-to-point navigation in 2D turbulent flows using reinforcement learning, Chaos 29, 103138 (2019).
- G. Reddy, A. Celani, T. J. Sejnowski and M. Vergassola, Learning to soar in turbulent environments, Proc. Natl. Acad. Sci. USA 113, 4877 (2016).
- G. Reddy, J. Wong-Ng, A. Celani, T. J. Sejnowski, and M. Vergassola, Glider soaring via reinforcement learning in the field, Nature (London) 562, 236 (2018).
- S. Verma, G. Novati, and P. Koumoutsakos, Efficient collective swimming by harnessing vortices through deep reinforcement learning, Proc. Natl. Acad. Sci. USA 115, 5849 (2018).
- C. David, J. Kennedy, and A. Ludlow, Finding of a sex pheromone source by gypsy moths released in the field, Nature (London) 303, 804 (1983).
- J. Murlis, J. S. Elkinton, and R. T. Carde, Odor plumes and how insects use them, Annu. Rev. Entomol. 37, 505 (1992).
- A. Loisy and C. Eloy, Searching for a source without gradients: How good is infotaxis and how to beat it, Proc. R. Soc. A 478, 20220118 (2022).
- K. V. B. Verano, E. Panizon, and A. Celani, Olfactory search with finite-state controllers, Proc. Natl. Acad. Sci. USA 120, e2304230120 (2023).
- W. Hanke, C. Brucker, and H. Bleckmann, The ageing of the low-frequency water disturbances caused by swimming goldfish and its possible relevance to prey detection, J. Exp. Biol. 203, 1193 (2000).
- K. Taira et al., Modal analysis of fluid flows: Applications and outlook, AIAA J. 58, 998 (2020).
- B. Colvert, K. K. Chen, and E. Kanso, Bioinspired sensory systems for shear flow detection, J. Nonlinear Sci. 27, 1183 (2017).
- F. van Breugel and M. H. Dickinson, Plume-tracking behavior of flying Drosophila emerges from a set of distinct sensory-motor reflexes, Curr. Biol. 24, 274 (2014).
- G. Reddy, B. I. Shraiman, and M. Vergassola, Sector search strategies for odor trail tracking, Proc. Natl. Acad. Sci. USA 119, e2107431118 (2022).
- T. Redaelli, C. Eloy, and E. Kanso, Efficient strategy to track a planktonic target from its hydrodynamic signature, in APS Division of Fluid Dynamics Meeting Abstracts (American Physical Society, Phoenix, Arizona, 2021), Q11-009.
- J. Liu, J. H. Costello, and E. Kanso, Flow physics of nutrient transport drives functional design of ciliates, Nat. Commun. 16, 4154 (2025).
- J. A. Farrell, J. Murlis, X. Long, W. Li, and R. T. Cardé, Filament-based atmospheric dispersion model to achieve short time-scale structure of odor plumes, Environ. Fluid Mech. 2, 143 (2002).
- IBAMR: An adaptive and distributed-memory parallel implementation of the immersed boundary (IB) method (2018), https://ibamr.github.io.
- A. P. S. Bhalla, R. Bale, B. E. Griffith, and N. A. Patankar, A unified mathematical framework and an adaptive numerical method for fluid–structure interaction with rigid, deforming, and elastic bodies, J. Comput. Phys. 250, 446 (2013).
- B. E. Griffith, R. D. Hornung, D. M. McQueen, and C. S. Peskin, An adaptive, formally second order accurate version of the immersed boundary method, J. Comput. Phys. 223, 10 (2007).
- G. T. Csanady, Turbulent Diffusion in the Environment (Springer Science and Business Media, New York, 2012).
- N. E. Leonard and E. Fiorelli, Virtual leaders, artificial potentials and coordinated control of groups, in Proceedings of the 40th IEEE Conference on Decision and Control (Cat. No. 01CH37228) (IEEE, Orlando, Florida, 2001), Vol. 3, pp. 2968–2973.
- J. Cochran and M. Krstic, Nonholonomic source seeking with tuning of angular velocity, Trans. Autom. Control, 54, 717 (2009).
- Y. Huang, J. Yen, and E. Kanso, Detection and tracking of chemical trails in bio-inspired sensory systems, Eur. J. Comput. Mech. 26, 98 (2017).
- L. Ristroph, J. C. Liao, and J. Zhang, Lateral line layout correlates with the differential hydrodynamic pressure on swimming fish, Phys. Rev. Lett. 114, 018102 (2015).
- B. Colvert and E. Kanso, Fishlike rheotaxis, J. Fluid Mech. 793, 656 (2016).
- J. Schulman, F. Wolski, P. Dhariwal, A. Radford, and O. Klimov, Proximal policy optimization algorithms, arXiv:1707.06347.
- Y. Jiao et al., Deep Dive into Model-free Reinforcement Learning for Biological and Robotic Systems: Theory and Practice, arXiv:2405.11457.
- See Supplemental Material at http://link.aps.org/supplemental/10.1103/qj9q-p7rg for Movies S1–S3 presents representative hydrodynamic trail-following trajectories for the RL-trained and RL-inspired control policies. It includes examples from the training domain as well as generalization to unseen flow conditions, spanning 2D and 3D wakes and turbulent plumes. Performance is further demonstrated across multiple sensing modalities, including flow speed, pressure, vorticity, and odor concentration.
- T. Seyde et al., Is bang-bang control all you need? solving continuous control with Bernoulli policies, Adv. Neural Inf. Proc. Syst. 34, 27209 (2021).
- V. Braitenberg, Vehicles: Experiments in Synthetic Psychology (MIT Press, Cambridge, Massachusetts, 1986).
- D. Shaikh and I. Rañó, Braitenberg vehicles as computational tools for research in neuro-science, Front. Bioeng. Biotechnol. 8, 565963 (2020).
- T. Salumäe, I. Rañó, O. Akanyeti, and M. Kruusmaa, Against the flow: A Braitenberg controller for a fish robot, in 2012 IEEE International Conference on Robotics and Automation (IEEE, Saint Paul, Minnesota, 2012), pp. 4210–4215.
- Y. Zhu, A. Nern, S. L. Zipursky, and M. A. Frye, Peripheral visual circuits functionally segregate motion and phototaxis behaviors in the fly, Curr. Biol. 19, 613 (2009).
- A. Lilienthal and T. Duckett, Experimental analysis of gas-sensitive Braitenberg vehicles, Adv. Robotics 18, 817 (2004).
- S. Heydari and E. Kanso, School cohesion, speed and efficiency are modulated by the swimmers flapping motion, J. Fluid Mech. 922, A27 (2021).
- G. Arranz, O. Flores, and M. Garcia-Villalba, Flow interaction of three-dimensional self propelled flexible plates in tandem, J. Fluid Mech. 931, A5 (2022).
- S. Heydari, H. Hang, and E. Kanso, Mapping spatial patterns to energetic benefits in groups of flow-coupled swimmers, elife 2024, 02 (2024).
- G. S. Triantafyllou, M. S. Triantafyllou, and M. A. Grosenbaugh, Optimal thrust development in oscillating foils with application to fish propulsion, J. Fluids Struct. 7, 205 (1993).
- D. Floryan, T. Van Buren, C. W. Rowley, and A. J. Smits, Scaling the propulsive performance of heaving and pitching foils, J. Fluid Mech. 822, 386 (2017).
- J. W. Newbolt, J. Zhang, and L. Ristroph, Flow interactions between uncoordinated flapping swimmers give rise to group cohesion, Proc. Natl. Acad. Sci. USA 116, 2419 (2019).
- P. Oteiza, I. Odstrcil, G. Lauder, R. Portugues, and F. Engert, A novel mechanism for mechanosensory-based rheotaxis in larval zebrafish, Nature (London) 547, 445 (2017).
- R. Venturelli et al., Hydrodynamic pressure sensing with an artificial lateral line in steady and unsteady flows, Bioinspiration Biomimetics 7, 036004 (2012).
- J.-H. Seo and R. Mittal, Improved swimming performance in schooling fish via leading-edge vortex enhancement, Bioinspiration Biomimetics 17, 066020 (2022).
- C. H. Williamson and A. Roshko, Vortex formation in the wake of an oscillating cylinder, J. Fluids Struct. 2, 355 (1988).
- T. Schnipper, A. Andersen, and T. Bohr, Vortex wakes of a flapping foil, J. Fluid Mech. 633, 411 (2009).
- S. Verma, C. Papadimitriou, N. Lüthen, G. Arampatzis, and P. Koumoutsakos, Optimal sensor placement for artificial swimmers, J. Fluid Mech. 884, A24 (2020).
- P. Weber et al., Optimal flow sensing for schooling swimmers, Biomimetics 5, 10 (2020).
- G. Vanwalleghem, K. Schuster, M. A. Taylor, I. A. Favre-Bulle, and E. K. Scott, Brain-wide mapping of water flow perception in zebrafish, J. Neurosci. 40, 4130 (2020).
- M. S. Madhav and N. J. Cowan, The synergy between neuroscience and control theory: The nervous system as inspiration for hard control challenges, Annu. Rev. Control Rob. Auton. Syst. 3, 243 (2020).
- R. A. Kastelein, P. J. Wensveen, J. M. Terhune, and C. A. de Jong, Near-threshold equal-loudness contours for harbor seals (Phoca vitulina) derived from reaction times during underwater audiometry: A preliminary study, J. Acoust. Soc. Am. 129, 488 (2011).
- P. Domenici and M. E. Hale, Escape responses of fish: A review of the diversity in motor control, kinematics and behaviour, J. Exp. Biol. 222, jeb166009 (2019).
- A. Roberts et al., The decision to move: response times, neuronal circuits and sensory memory in a simple vertebrate, Proc. R. Soc. B 286, 20190297 (2019).
- P. Domenici and R. W. Blake, The kinematics and performance of fish fast-start swimming, J. Exp. Biol. 200, 1165 (1997).
- D. Weihs, The mechanism of rapid starting of slender fish, Biorheology 10, 343 (1973).
- K. Sato et al., Stroke frequency, but not swimming speed, is related to body size in free-ranging seabirds, pinnipeds and cetaceans, Proc. R. Soc. B 274, 471 (2007).
- J. Brett, The relation of size to rate of oxygen consumption and sustained swimming speed of sockeye salmon (Oncorhynchus nerka), J. Fish. Board Can. 22, 1491 (1965).
- M. Kasapi, P. Domenici, R. Blake, and D. Harper, The kinematics and performance of escape responses of the knifefish Xenomystus nigri, Can. J. Zool. 71, 189 (1993).
- P. Domenici and R. W. Blake, The kinematics and performance of the escape response in the angelfish (Pterophyllum eimekei), J. Exp. Biol. 156, 187 (1991).
- D. G. Harper and R. W. Blake, Fast-start performance of rainbow trout Salmo gairdneri and northern pike Esox lucius, J. Exp. Biol. 150, 321 (1990).
- S. J. Peake and A. P. Farrell, Locomotory behaviour and post-exercise physiology in relation to swimming speed, gait transition and metabolism in free-swimming smallmouth bass (Micropterus dolomieu), J. Exp. Biol. 207, 1563 (2004).
- J. D. Stieglitz, E. M. Mager, R. H. Hoenig, D. D. Benetti, and M. Grosell, Impacts of deepwater horizon crude oil exposure on adult mahi-mahi (Coryphaena hippurus) swim performance, Environ. Toxicol. Chem. 35, 2613 (2016).
- J. D. Kieffer, D. Alsop, and C. M. Wood, A respirometric analysis of fuel use during aerobic swimming at different temperatures in rainbow trout (Oncorhynchus mykiss), J. Exp. Biol. 201, 3123 (1998).
- N. C. Wegner, M. A. Drawbridge, and J. R. Hyde, Reduced swimming and metabolic fitness of aquaculture-reared California yellowtail (Seriola dorsalis) in comparison to wild-caught conspecifics, Aquaculture 486, 51 (2018).
- B. Block et al., Environmental preferences of yellowfin tuna (Thunnus albacares) at the northern extent of its range, Mar. Biol. 130, 119 (1997).
- P. W. Webb and R. S. Keyes, Division of labor between median fins in swimming dolphin (Pisces: Coryphaenidae), Copeia 1981, 901 (1981).
- F. E. Fish, J. Hurley, and D. P. Costa, Maneuverability by the sea lion Zalophus californianus: Turning performance of an unstable body design, J. Exp. Biol. 206, 667 (2003).
- Some of these data were available in the min-max range (e.g., [102, 103]), which we converted to mean and standard deviation using the range rule [120].
- G. Van Belle, Statistical Rules of Thumb (John Wiley & Sons, Hoboken, New Jersey, 2011).
- Y. Jiao, B. Colvert, Y. Man, M. J. McHenry, and E. Kanso, Evaluating evasion strategies in zebrafish larvae, Proc. Natl. Acad. Sci. USA 120, e2218909120 (2023).
- S. A. Burden, T. Libby, K. Jayaram, S. Sponberg, and J. M. Donelan, Why animals can outrun robots, Sci. Rob. 9, eadi9754 (2024).
- Y. Jiao et al., Learning to swim in potential flow, Phys. Rev. Fluids 6, 050505 (2021).
- J. Yu, L. Liu, L. Wang, M. Tan, and D. Xu, Turning control of a multilink biomimetic robotic fish, IEEE Trans. Rob. 24, 201 (2008).
- K. Hirata, T. Takimoto, and K. Tamura, Study on turning performance of a fish robot, in First International Symposium on Aqua Bio-Mechanisms (Society of Aero-Aqua Bio-Mechanism, Honolulu, Hawaii, 2000), pp. 287–292.
- J. Liang, T. Wang, and L. Wen, Development of a two-joint robotic fish for real-world exploration, J. Field Rob. 28, 70 (2011).
- C. S. Peskin, Numerical analysis of blood flow in the heart, J. Comput. Phys. 25, 220 (1977).
- R. Mittal et al., A versatile sharp interface immersed boundary method for incompressible flows with complex boundaries, J. Comput. Phys. 227, 4825 (2008).
- A. Vargas, R. Mittal, and H. Dong, A computational study of the aerodynamic performance of a dragonfly wing section in gliding flight, Bioinspiration Biomimetics 3, 026004 (2008).
- R. Mittal and G. Iaccarino, Immersed boundary methods, Annu. Rev. Fluid Mech. 37, 239 (2005).
- OpenAI Spinning up documentation—Proximal policy optimization (2018), https://spinningup.openai.com/en/latest/algorithms/ppo.html.
- OpenAI Github baseline—Proximal policy optimization (2017), https://github.com/openai/baselines/tree/master/baselines/ppo2.