Jaume Casademunt

• Professor of Physics
• University of Barcelona

Active fluids display spontaneous turbulent-like flows known as active turbulence. Recent work revealed that these flows have universal features, independent of the material properties and of the presence of topological defects. However, the differences between defect-laden and defect-free active turbulence remain largely unexplored. Here, by means...

Active nematics exhibit spontaneous flows through a well-known linear instability of the uniformly-aligned quiescent state. Here we show that even a linearly stable uniform state can experience a nonlinear instability, resulting in a discontinuous transition to spontaneous flows. In this case, quiescent and flowing states may coexist. Through a wea...

The efficient transport of fluids through disordered media requires a thorough understanding of how the driving rate affects two-phase interface propagation. Despite our understanding of front dynamics in homogeneous environments, as well as how medium heterogeneities shape fluid interfaces at rest, little is known about the effects of localized to...

The directed migration of cellular clusters enables morphogenesis, wound healing and collective cancer invasion. Gradients of substrate stiffness direct the migration of cellular clusters in a process called collective durotaxis, but the underlying mechanisms remain unclear. Here we unveil a connection between collective durotaxis and the wetting p...

The directed migration of cellular clusters enables morphogenesis, wound healing, and collective cancer invasion. Gradients of substrate stiffness are known to direct the migration of cellular clusters in a process called collective durotaxis, but underlying mechanisms remain unclear. Here, we unveil a connection between collective durotaxis and th...

Active fluids exhibit spontaneous flows with complex spatiotemporal structure, which have been observed in bacterial suspensions, sperm cells, cytoskeletal suspensions, self-propelled colloids, and cell tissues. Despite occurring in the absence of inertia, chaotic active flows are reminiscent of inertial turbulence, and hence they are known as acti...

Many types of motile cells perform durotaxis, namely directed migration following gradients of substrate stiffness. Recent experiments have revealed that cell monolayers can migrate toward stiffer regions even when individual cells do not—a phenomenon known as collective durotaxis. Here, we address the spontaneous motion of finite cohesive cell mon...

Many types of motile cells perform durotaxis, namely, directed migration following gradients of substrate stiffness. Recent experiments have revealed that cell monolayers can migrate toward stiffer regions even when individual cells do not -- a phenomenon known as collective durotaxis. Here we address the spontaneous motion of finite cohesive cell...

Active fluids exhibit complex turbulentlike flows at low Reynolds number. Recent work predicted that 2D active nematic turbulence follows scaling laws with universal exponents. However, experimentally testing these predictions is conditioned by the coupling to the 3D environment. Here, we measure the spectrum of the kinetic energy E(q) in an active...

Active fluids exhibit spontaneous flows with complex spatiotemporal structure, which have been observed in bacterial suspensions, sperm cells, cytoskeletal suspensions, self-propelled colloids, and cell tissues. Despite occurring in the absence of inertia, chaotic active flows are reminiscent of inertial turbulence, and hence they are known as acti...

Recent experiments have shown that the spontaneous activity of developing dissociated neuronal cultures can be described as a process of highly inhomogeneous nucleation and front propagation due to the localization of noise activity, i.e. , noise focusing. However, the basic understanding of the mechanisms of noise build-up leading to the nucleatio...

Active fluids exhibit complex turbulent-like flows at low Reynolds number. Recent work predicted that 2d active nematic turbulence follows universal scaling laws. However, experimentally testing these predictions is conditioned by the coupling to the 3d environment. Here, we measure the spectrum of the kinetic energy, E(q), in an active nematic fil...

Recent experiments have shown that the spontaneous activity of young dissociated neuronal cultures can be described as a process of highly inhomogeneous nucleation and front propagation due to the localization of noise activity, i.e., noise focusing. However, the basic understanding of the mechanisms of noise build-up leading to the nucleation rema...

We design, construct, and test a device for controlled boiling by means of local vapor bubble formation in a cavity that injects vapor bubbles in a capillary crossflow. The outcome of the device is a regular slug flow of approximately monodisperse bubbles that can be used for a variety of applications. The setup is designed to be independent of gra...

A landmark of turbulence is the emergence of universal scaling laws, such as Kolmogorov’s E(q) ~ q−5∕3 scaling of the kinetic energy spectrum of inertial turbulence with the wavevector q. In recent years, active fluids have been shown to exhibit turbulent-like flows at low Reynolds number. However, the existence of universal scaling properties in t...

Living tissues undergo wetting transitions: On a surface, they can either form a droplet-like cell aggregate or spread as a monolayer of migrating cells. Tissue wetting depends not only on the chemical but also on the mechanical properties of the substrate. Here, we study the role of substrate stiffness in tissue spreading, which we describe by mea...

A landmark of turbulence is the emergence of universal scaling laws, such as Kolmogorov’s E(q) ~ q−5∕3 scaling of the kinetic energy spectrum of inertial turbulence with the wavevector q. In recent years, active fluids have been shown to exhibit turbulent-like flows at low Reynolds number. However, the existence of universal scaling properties in t...

We introduce a minimal hydrodynamic model of polarization, migration, and deformation of a biological cell confined between two parallel surfaces. In our model, the cell is driven out of equilibrium by an active cytsokeleton force that acts on the membrane. The cell cytoplasm, described as a viscous droplet in the Darcy flow regime, contains a diff...

Brownian ratchets are shown to feature a nontrivial vanishing-noise limit where the dynamics is reduced to a stochastic alternation between two deterministic circle maps (quasi-deterministic ratchets). Motivated by cooperative dynamics of molecular motors, here we solve exactly the problem of two interacting quasi-deterministic ratchets. We show th...

An amendment to this paper has been published and can be accessed via a link at the top of the paper.

A landmark of turbulence is the emergence of universal scaling laws, such as Kolmogorov's E(q)∼q^{−5/3} scaling of the kinetic energy spectrum of inertial turbulence with the wave vector q. In recent years, active fluids have been shown to exhibit turbulent-like flows at low Reynolds number. However, the existence of universal scaling properties in...

Living tissues undergo wetting transitions: On a surface, they can either form a droplet-like cell aggregate or spread as a monolayer of migrating cells. Tissue wetting depends not only on the chemical but also on the mechanical properties of the substrate. Here, we study the role of substrate stiffness in tissue spreading, which we describe by mea...

A landmark of turbulence is the emergence of universal scaling laws, such as Kolmogorov's $E(q)\sim q^{-5/3}$ scaling of the kinetic energy spectrum of inertial turbulence with the wave vector $q$. In recent years, active fluids have been shown to exhibit turbulent-like flows at low Reynolds number. However, the existence of universal scaling prope...

The kinesin-3 motor KIF1A is involved in long-ranged axonal transport in neurons. In order to ensure vesicular delivery, motors need to navigate the microtubule lattice and overcome possible roadblocks along the way. The single-headed form of KIF1A is a highly diffusive motor that has been shown to be a prototype of Brownian motor by virtue of a we...

During the spreading of epithelial tissues, the advancing tissue front often develops fingerlike protrusions. Their resemblance to traditional viscous fingering patterns in driven fluids suggests that epithelial fingers could arise from an interfacial instability. However, the existence and physical mechanism of such a putative instability remain u...

During the spreading of epithelial tissues, the advancing tissue front often develops fingerlike protrusions. Their resemblance to traditional viscous fingering patterns in driven fluids suggests that epithelial fingers could arise from an interfacial instability. However, the existence and physical mechanism of such a putative instability remain u...

Development, regeneration and cancer involve drastic transitions in tissue morphology. In analogy with the behavior of inert fluids, some of these transitions have been interpreted as wetting transitions. The validity and scope of this analogy are unclear, however, because the active cellular forces that drive tissue wetting have been neither measu...

Active turbulence describes a flow regime that is erratic, and yet endowed with a characteristic length scale¹. It arises in animate soft-matter systems as diverse as bacterial baths², cell tissues³ and reconstituted cytoskeletal preparations⁴. However, the way that these turbulent dynamics emerge in active systems has so far evaded experimental sc...

The kinesin-3 motor KIF1A is involved in long-ranged axonal transport in neurons. In order to ensure vesicular delivery, motors need to navigate the microtubule lattice and overcome possible roadblocks along the way. The single-headed form of KIF1A is a highly diffusive motor that has been shown to be a prototype of Brownian motor by virtue of a we...

Development, regeneration and cancer involve drastic transitions in tissue morphology. In analogy with the behaviour of inert fluids, some of these transitions have been interpreted as wetting transitions. The validity and scope of this analogy are unclear, however, because the active cellular forces that drive tissue wetting have been neither meas...

Lattice-Boltzmann simulations of a turbulent duct flow have been carried out to obtain trajectories of passive tracers in the conditions of a series of microgravity experiments of turbulent bubble suspensions. The statistics of these passive tracers are compared to the corresponding measurements for single-bubble and bubble-pair statistics obtained...

A new experimental setup for the generation of homogeneous, monodisperse bubble suspensions in turbulent duct flows in microgravity has been designed and tested in drop tower experiments. The setup provides independent control of bubble size, void fraction and degree of turbulence. The device combines several slug-flow injectors that produce monodi...

Mixed-order phase transitions display a discontinuity in the order parameter like first-order transitions yet feature critical behavior like second-order transitions. Such transitions have been predicted for a broad range of equilibrium and nonequilibrium systems, but their experimental observation has remained elusive. Here, we analytically predic...

Significance Phase transitions are usually classified in two categories: First-order transitions are discontinuous, and second-order transitions are continuous and exhibit critical behavior. However, the so-called mixed-order transitions combine features of both types, such as being discontinuous yet featuring a diverging correlation length. Such t...

We consider the displacement of an inviscid fluid (air) by a viscous fluid (oil) in a narrow channel with gap-thickness modulations. The interfacial dynamics of this problem is strongly nonlocal and exhibits competing effects from capillarity and permeability. We derive analytical predictions of steady-state front morphologies, which are exact at l...

We introduce a coarse-grained stochastic model for the spontaneous activity of neuronal cultures to explain the phenomenon of noise focusing, which entails localization of the noise activity in excitable networks with metric correlations. The system is modeled as a continuum excitable medium with a state-dependent spatial coupling that accounts for...

We present a hydrodynamic model of spreading epithelial monolayers as polar viscous fluids, with active contractility and traction on the substrate. The combination of both active forces generate an instability that leads to nonlinear traveling waves, which propagate in the direction of polarity with characteristic time scales that depend on contac...

We present a hydrodynamic model of spreading epithelial monolayers as polar viscous fluids, with active contractility and traction on the substrate. The combination of both active forces generate an instability that leads to nonlinear traveling waves, which propagate in the direction of polarity with characteristic time scales that depend on contac...

The physical basis of flagellar and ciliary beating is a major problem in biology which is still far from completely understood. The fundamental cytoskeleton structure of cilia and flagella is the axoneme, a cylindrical array of microtubule doublets connected by passive crosslinkers and dynein motor proteins. The complex interplay of these elements...

We derive the constitutive equations of an active polar gel from a model for the dynamics of elastic molecules that link polar elements. Molecular binding kinetics induces the fluidization of the material, giving rise to Maxwell viscoelasticity and, provided that detailed balance is broken, to the generation of active stresses. We give explicit exp...

We derive the constitutive equations of an active polar gel from a model for the dynamics of elastic molecules that link polar elements. Molecular binding kinetics induces the fluidization of the material, giving rise to Maxwell viscoelasticity and, provided that detailed balance is broken, to the generation of active stresses. We give explicit exp...

Collective cell migration in spreading epithelia in controlled environments has become a landmark in our current understanding of fundamental biophysical processes in development, regeneration, wound healing or cancer. Epithelial monolayers are treated as thin layers of a viscous fluid that exert active traction forces on the substrate. The model i...

We derive the constitutive equations of an active polar gel from a model for the dynamics of elastic molecules that link polar elements. Molecular binding kinetics induces the fluidization of the material, giving rise to Maxwell viscoelasticity and, provided that detailed balance is broken, to the generation of active stresses. We give explicit exp...

Metastable phases may be spontaneously formed from other metastable phases through nucleation. Here we demonstrate the spontaneous formation of a metastable phase from an unstable equilibrium by spinodal decomposition, which leads to a transient coexistence of stable and metastable phases. This phenomenon is generic within the recently introduced s...

Metastable phases may be spontaneously formed from other metastable phases through nucleation. Here we demonstrate the spontaneous formation of a metastable phase from an unstable equilibrium by spinodal decomposition, which leads to a transient coexistence of stable and metastable phases. This phenomenon is generic within the recently introduced s...

Illustration of the formation of the least energetic interface when possible, even when the most energetic interface is initially present.

Supplementary Figure 1, Supplementary Note 1, Supplementary Discussion, Supplementary Methods and Supplementary References

Illustration of the coexistence of two equilibrium phases connected by two physically different interfaces.

Illustration of the formation of metastable domains by spinodal decomposition and their subsequent elimination by front propagation.

Metastable phases may be spontaneously formed from other metastable phases through nucleation. Here we demonstrate the spontaneous formation of a metastable phase from an unstable equilibrium by spinodal decomposition, which leads to a transient coexistence of stable and metastable phases. This phenomenon is generic within the recently introduced s...

The physical basis of flagellar and ciliary beating is a major problem in biology which is still far from completely understood. The fundamental cytoskeleton structure of cilia and flagella is the axoneme, a cylindrical array of microtubule doublets connected by passive crosslinkers and dynein motor proteins. The complex interplay of these elements...

We study the nucleation of blebs, i.e., protrusions arising from a local detachment of the membrane from the cortex of a cell. Based on a simple model of elastic linkers with force-dependent kinetics, we show that bleb nucleation is governed by membrane peeling. By this mechanism, the growth or shrinkage of a detached membrane patch is completely d...

We study the nucleation of blebs, i.e., protrusions arising from a local detachment of the membrane from the cortex of a cell. Based on a simple model of elastic linkers with force-dependent kinetics, we show that bleb nucleation is governed by membrane peeling. By this mechanism, the growth or shrinkage of a detached membrane patch is completely d...

We study the nucleation of blebs, i.e., protrusions arising from a local detachment of the membrane from the cortex of a cell. Based on a simple model of elastic linkers with force-dependent kinetics, we show that bleb nucleation is governed by membrane peeling. By this mechanism, the growth or shrinkage of a detached membrane patch is completely d...

We propose a model for membrane-cortex adhesion which couples membrane deformations, hydrodynamics and kinetics of membrane-cortex ligands. In its simplest form, the model gives explicit predictions for the critical pressure for membrane detachment and for the value of adhesion energy. We show that these quantities exhibit a significant dependence...

We study the 2D crystalline phases of paramagnetic colloidal particles with dipolar interactions and constrained on a periodic substrate. Combining theory, simulation, and experiments we demonstrate a new scenario of first-order phase transitions that occurs via a complete inversion of the energy landscape, featuring non-conventional properties tha...

The kinesin-3 motor KIF1A is in charge of vesicular transport in neuronal axons. Its single-headed form is known to be very inefficient due to the presence of a diffusive state in the mechanochemical cycle. However, recent theoretical studies have suggested that these motors could largely enhance force generation by working in teams. Here we test t...

A detailed statistical analysis of bubble dispersion in turbulent jets based on data from drop tower experiments is presented here. A stochastic model is also introduced in order to capture these statistics to a large extent, treating bubbles as passive tracers with a local diffusivity given by a k- ε description of the turbulence. Bubble-bubble an...

We propose a model for membrane-cortex adhesion that couples membrane deformations, hydrodynamics, and kinetics of membrane-cortex ligands. In its simplest form, the model gives explicit predictions for the critical pressure for membrane detachment and for the value of adhesion energy. We show that these quantities exhibit a significant dependence...

We propose a model for membrane-cortex adhesion that couples membrane deformations, hydrodynamics, and kinetics of membrane-cortex ligands. In its simplest form, the model gives explicit predictions for the critical pressure for membrane detachment and for the value of adhesion energy. We show that these quantities exhibit a significant dependence...

We study the 2D crystalline phases of paramagnetic colloidal particles with dipolar interactions and constrained on a periodic substrate. Combining theory, simulation, and experiments, we demonstrate a new scenario of first-order phase transitions that occurs via a complete inversion of the energy landscape, featuring nonconventional properties tha...

We study the 2D crystalline phases of paramagnetic colloidal particles with dipolar interactions and constrained on a periodic substrate. Combining theory, simulation, and experiments, we demonstrate a new scenario of first-order phase transitions that occurs via a complete inversion of the energy landscape, featuring nonconventional properties tha...

We perform a characterization of a recently reported microbubble injector in conditions relevant to microgravity. Injection of bubbles is based on the generation of a slug flow in a capillary T-junction, whose operation is robust to changes in the gravity level. We address questions regarding the performance under different working regimes. In part...

We study a generic two-state model for an assembly of molecular motors which is described by means of a pair of integro-partial differential equations and leads to oscillatory motion in the presence of an elastic coupling to its environment. We discuss a reduction of the system to a minimal set of three ordinary differential equations that successf...

We study and classify the time-dependent morphologies of polarized tissues subject to anisotropic but spatially homogeneous growth. Extending previous studies, we model the tissue as a fluid, and discuss the interplay of the active stresses generated by the anisotropic cell division and three types of passive mechanical forces: viscous stresses, fr...

Cultured neurons in vitro quickly connect to one another to establish a spontaneously active network within a week. The resulting neuronal network is characterized by a combination of excitatory and inhibitory integrate–and–fire units coupled through synaptic connections , and that interact in a highly nonlinear manner. The nonlinear behavior emerg...

We study in detail the cooperative action of small groups of KIF1A motors in its monomeric (single-headed) form within an arrangement relevant to vesicle traffic or membrane tube extraction. It has been recently shown that under these circumstances, the presence of a finite dwell time in the motor cycle contributes to remarkably enhance collective...

At early stages of development, neuronal cultures in vitro spontaneously reach a coherent state of collective firing in a pattern of nearly periodic global bursts. Although understanding the spontaneous activity of neuronal networks is of chief importance in neuroscience, the origin and nature of that pulsation has remained elusive. By combining hi...

KIF1A is a kinesin motor protein that can work processively in a monomeric (single-headed) form by using a noise-driven ratchet mechanism. Here, we show that the combination of a passive diffusive state and finite-time kinetics of adenosine triphosphate hydrolysis provides a powerful mechanism of cooperative force generation, implying for instance...

We show that actin lamellar fragments driven solely by polymerization forces at the bounding membrane are generically motile when the circular symmetry is spontaneously broken, with no need of molecular motors or global polarization. We base our study on a nonlinear analysis of a recently introduced minimal model [Callan-Jones et al., Phys. Rev. Le...

The residual accelerations that are typically present in microgravity environments (g-jitters) contain a broad spectrum of frequencies and may be modeled as stochastic processes. Their effects on the quality of the semiconductor crystals are analyzed here quantitatively with direct numerical simulation. In particular we focus on the dopant segregat...

We analyze the dynamics of overlapping antiparallel treadmilling microtubules in the presence of crosslinking processive motor proteins that counterbalance an external force. We show that coupling the force-dependent velocity of motors and the kinetics of motor exchange with a bath in the presence of treadmilling leads generically to oscillatory be...

In vitro neuronal networks of dissociated hippocampal or cortical tissues are one of the most attractive model systems for the physics and neuroscience communities. Cultured neurons grow and mature, develop axons and dendrites, and quickly connect to their neighbors to establish a spontaneously active network within a week. The resulting neuronal...

We discuss the use of phase-field methods in problems of interface dynamics where topological singularities are relevant. We illustrate our discussion on the case study of centrifugally driven Hele-Shaw flows, where the proposed analysis has been recently tested. We argue that the phase-field formulation of a given sharp-interface model is not only...

Position of the division events and actual duration of the cell cycle. When a cell divides, the position of the mother cell is marked by a circle just before the cleavage event. The color code indicates the actual duration of the cell cycle. In average, the actual duration of the cell cycle is hours for cells at the compartment bulk and hours for c...

In silico laser ablation of the edge of a cell with a bulk character. Left: cells with an ablated edge appear in red. Right: separation of the vertexes that correspond to the ablated edge as a function of time. (AVI)

The actomyosin cable removal produces a DV organizer that lacks robustness. While in this video there is not cell intermingling the organizer is only one cell wide at different locations and threatens to break. (AVI)

The Hertwig rule is required for keeping cell segregation. If no rule for cell cleavage is prescribed the DV organizer breaks. Here the cleavage direction is set at random. (AVI)

Dynamics of DV organizer (red) in an in silico wild-type wing imaginal disc. (AVI)

A clone of rapid duplicating cells (blue) exerts a differential pressure over the DV organizer (red) that bends. Still, under wild-type conditions, the organizer remains intact and restricts the cellular lineages. (AVI)

An ectopic organizer (red) evolves minimizing the surface energy. The organizer is initially set with an eight-shaped conformation and ends with a circular one. (AVI)

The removal of the organizer produces cell mix. In the absence of an organizer, cells from opposite compartments do not respect the DV boundary and start to intermingle. (AVI)

(A) “Toy” regulatory network: Gene-protein B is positively regulated by gene-protein A. On the other hand, this interaction is negatively regulated by gene-protein C. Hill functions, with a given degree of cooperativity, β, are assumed to effectively model gene-protein regulation. (B) Hill functions for distinct values of cooperativity. As β increa...

Correlations in the cell cycle phase. Here all cells have a bulk character and the initial phase of each cell is taken at random. The color code indicates the progression of the cell cycle: redyellowgreenblue. A cell and its progeny maintain the coherence and synchronization of the cell cycle phases during a temporal window. Yet, in the long term,...

Differential affinities are required for a robust development of the organizer. If all cells have the same affinity then the DV organizer easily breaks. (AVI)

Effect of the cell cycle duration. If there is not a differential cell cycle duration of the cells at the DV organizer with respect to cells at the bulk, the organizer becomes wider and the boundary wiggly. (AVI)

We analyse the dynamics of overlapping antiparallel treadmilling microtubules in the presence of crosslinking processive motor proteins that counterbalance an external force. We show that coupling the force-dependent velocity of motors and the kinetics of motor exchange with a bath in the presence of treadmilling leads generically to oscillatory be...

Shaping the primordia during development relies on forces and mechanisms able to control cell segregation. In the imaginal discs of Drosophila the cellular populations that will give rise to the dorsal and ventral parts on the wing blade are segregated and do not intermingle. A cellular population that becomes specified by the boundary of the dorsa...

We study the cooperative dynamics of Brownian motors moving along a one-dimensional track when an external load is applied to the leading motor, mimicking molecular motors pulling on membrane-bound cargoes in intracellular traffic. Due to the asymmetric loading, self-organized motor clusters form spontaneously. We model the motors with a two-state...

Bleb-based cell motility proceeds by the successive inflation and retraction of large spherical membrane protrusions ("blebs") coupled with substrate adhesion. In addition to their role in moti-lity, cellular blebs constitute a remarkable illustration of the dyna-mical interactions between the cytoskeletal cortex and the plasma membrane. Here we st...

Bleb-based cell motility proceeds by the successive inflation and retraction of large spherical membrane protrusions ("blebs") coupled with substrate adhesion. In addition to their role in motility, cellular blebs constitute a remarkable illustration of the dynamical interactions between the cytoskeletal cortex and the plasma membrane. Here we stud...