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Cilia and flagella often exhibit synchronized behavior; this includes phase locking, as seen in {\it Chlamydomonas}, and metachronal wave formation in the respiratory cilia of higher organisms. Since the observations by Gray and Rothschild…

Soft Condensed Matter · Physics 2016-05-19 Kirsty Y. Wan , Raymond E. Goldstein

Despite evidence for a hydrodynamic origin of flagellar synchronization between different eukaryotic cells, recent experiments have shown that in single multi-flagellated organisms, coordination hinges instead on direct basal body…

Biological Physics · Physics 2018-10-12 Yujie Liu , Rory Claydon , Marco Polin , Douglas R. Brumley

Microscale fluid flows generated by ensembles of beating eukaryotic flagella are crucial to fundamental processes such as development, motility and sensing. Despite significant experimental and theoretical progress, the underlying physical…

Soft Condensed Matter · Physics 2014-03-11 Douglas R. Brumley , Kirsty Y. Wan , Marco Polin , Raymond E. Goldstein

Beating flagella exhibit a variety of synchronization modes. This synchrony has long been attributed to hydrodynamic coupling between the flagella. However, recent work with flagellated algae indicates that a mechanism internal to the cell,…

Biological Physics · Physics 2020-11-18 Hanliang Guo , Yi Man , Kirsty Y. Wan , Eva Kanso

In a multitude of life's processes, cilia and flagella are found indispensable. Recently, the biflagellated chlorophyte alga Chlamydomonas has become a model organism for the study of ciliary coordination and synchronization. Here, we use…

Biological Physics · Physics 2014-02-28 Kirsty Y. Wan , Kyriacos C. Leptos , Raymond E. Goldstein

The present habilitation thesis in theoretical biological physics addresses two central dynamical processes in cells and organisms: (i) active motility and motility control and (ii) self-organized pattern formation. The unifying theme is…

Cell Behavior · Quantitative Biology 2018-03-21 Benjamin M. Friedrich

The eukaryotic flagellum beats with apparently unfailing periodicity, yet responds rapidly to stimuli. Like the human heartbeat, flagellar oscillations are now known to be noisy. Using the alga \textit{C. reinhardtii}, we explore three…

Soft Condensed Matter · Physics 2014-12-03 Kirsty Y. Wan , Raymond E. Goldstein

Cilia and flagella exhibit regular bending waves that perform mechanical work on the surrounding fluid, to propel cellular swimmers and pump fluids inside organisms. Here, we quantify a force-velocity relationship of the beating flagellum,…

Cell Behavior · Quantitative Biology 2016-12-21 Gary S. Klindt , Christian Ruloff , Christian Wanger , Benjamin M. Friedrich

The unicellular green algae Chlamydomonas swims with two flagella, which can synchronize their beat. Synchronized beating is required to swim both fast and straight. A long-standing hypothesis proposes that synchronization of flagella…

Cell Behavior · Quantitative Biology 2013-11-26 Veikko Geyer , Frank Jülicher , Jonathon Howard , Benjamin M Friedrich

Groups of beating flagella or cilia often synchronize so that neighboring filaments have identical frequencies and phases. A prime example is provided by the unicellular biflagellate Chlamydomonas reinhardtii, which typically displays…

Sperm are propelled by bending waves travelling along the flagellum. During steering in gradients of sensory cues, sperm adjust the flagellar beat waveform. Symmetric and asymmetric beat waveforms produce straight and curved swimming paths,…

It is now well established that nearby beating pairs of eukaryotic flagella or cilia typically synchronize in phase. A substantial body of evidence supports the hypothesis that hydrodynamic coupling between the active filaments, combined…

Soft Condensed Matter · Physics 2016-10-12 Raymond E. Goldstein , Eric Lauga , Adriana I. Pesci , Michael R. E. Proctor

Many biological microorganisms and artificial microswimmers react to external cues of environmental gradients by changing their swimming directions. We study here the behavior of eukarytic flagellated microswimmers in linear viscosity…

Soft Condensed Matter · Physics 2026-05-29 Shubham Anand , Jens Elgeti , Gerhard Gompper

The motility of microorganisms is influenced greatly by their hydrodynamic interactions with the fluidic environment they inhabit. We show by direct experimental observation of the bi-flagellated alga Chlamydomonas reinhardtii that fluid…

Biological Physics · Physics 2015-11-03 Boyang Qin , Arvind Gopinath , Jing Yang , Jerry P Gollub , Paulo E Arratia

Living creatures exhibit a remarkable diversity of locomotion mechanisms, evolving structures specialised for interacting with their environment. In the vast majority of cases, locomotor behaviours such as flying, crawling, and running, are…

Biological Physics · Physics 2020-07-28 Kirsty Y. Wan

Synchronisation is often observed in the swimming of flagellated cells, either for multiple appendages on the same organism or between the flagella of nearby cells. Beating cilia are also seen to synchronise their dynamics. In 1951, Taylor…

Biological Physics · Physics 2021-04-28 Weida Liao , Eric Lauga

The eukaryotic flagellum beats periodically, driven by the oscillatory dynamics of molecular motors, to propel cells and pump fluids. Small, but perceivable fluctuations in the beat of individual flagella have physiological implications for…

Cell Behavior · Quantitative Biology 2015-06-18 Rui Ma , Gary S. Klindt , Ingmar H. Riedel-Kruse , Frank Jülicher , Benjamin M. Friedrich

While hydrodynamic coupling has long been considered essential for synchronisation of eukaryotic flagella, recent experiments on the unicellular biflagellate model organism {\it Chlamydomonas} demonstrate that -- at the single cell level --…

Biological Physics · Physics 2025-04-04 Luc Zorrilla , Antoine Allard , Krish Desai , Marco Polin

Sperm modulate their flagellar symmetry to navigate through complex physico-chemical environments and achieve reproductive function. Yet it remains elusive how sperm swim forwards despite the inherent asymmetry of several components that…

Biological Physics · Physics 2023-07-21 Xiaomeng Ren , Hermes Bloomfield-Gadêlha

Cellular appendages such as cilia and flagella represent universal tools enabling cells and microbes, among other essential functionalities, to propel themselves in diverse environments. In its planktonic, i.e. freely swimming, state the…

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