@phdthesis{21393, abstract = {This thesis documents a voyage towards truth and beauty via formal verification of theorems. To this end, we develop libraries in Lean 4 that present definitions and results from diverse areas of MathematiCS (i.e., Mathematics and Computer Science). The aim is to create code that is understandable, believable, useful, and elegant. The code should stand for itself as much as possible without a need for documentation; however, this text redundantly documents our code artifacts and provides additional context that isn’t present in the code. This thesis is written for readers who know Lean 4 but are not familiar with any of the topics presented. We manifest truth and beauty in three formalized areas of MathematiCS. We formalize general grammars in Lean 4 and use grammars to show closure of the class of type-0 languages under four operations; union, reversal, concatenation, and the Kleene star. Our second stop is the theory of optimization. Farkas established that a system of linear inequalities has a solution if and only if we cannot obtain a contradiction by taking a linear combination of the inequalities. We state and formally prove several Farkas-like theorems over linearly ordered fields in Lean 4. Furthermore, we extend duality theory to the case when some coefficients are allowed to take “infinite values”. Additionally, we develop the basics of the theory of optimization in terms of the framework called General-Valued Constraint Satisfaction Problems, and we prove that, if a Rational-Valued Constraint Satisfaction Problem template has symmetric fractional polymorphisms of all arities, then its basic LP relaxation is tight. Our third stop is matroid theory. Seymour’s decomposition theorem is a hallmark result in matroid theory, presenting a structural characterization of the class of regular matroids. We aim to formally verify Seymour’s theorem in Lean 4. First, we build a library for working with totally unimodular matrices. We define binary matroids and their standard representations, and we prove that they form a matroid in the sense how Mathlib defines matroids. We define regular matroids to be matroids for which there exists a full representation rational matrix that is totally unimodular, and we prove that all regular matroids are binary. We define 1-sum, 2-sum, and 3 sum of binary matroids as specific ways to compose their standard representation matrices. We prove that the 1-sum, the 2-sum, and the 3-sum of regular matroids are a regular matroid, which concludes the composition direction of the Seymour’s theorem. The (more difficult) decomposition direction remains unproved. In the pursuit of truth, we focus on identifying the trusted code in each project and presenting it faithfully. We emphasize the readability and believability of definitions rather than choosing definitions that are easier to work with. In search for beauty, we focus on the philosophical framework of Roger Scruton, who emphasizes that beauty is not a mere decoration but, most importantly, beauty is the means for shaping our place in the world and a source of redemption, where it can be viewed as a substitute for religion.}, author = {Dvorak, Martin}, isbn = {978-3-99078-074-9}, issn = {2663-337X}, pages = {160}, publisher = {Institute of Science and Technology Austria}, title = {{Pursuit of truth and beauty in Lean 4 : Formally verified theory of grammars, optimization, matroids}}, doi = {10.15479/AT-ISTA-21393}, year = {2026}, } @article{21501, abstract = {Kinetically constrained models were originally introduced to capture slow relaxation in glassy systems, where dynamics are hindered by local constraints instead of energy barriers. Their quantum counterparts have recently drawn attention for exhibiting highly degenerate eigenstates at zero energy—known as zero modes—stemming from chiral symmetry. Yet, the structure and implications of these zero modes remain poorly understood. In this work, we focus on the properties of the zero mode subspace in quantum kinetically constrained models with a U(1) particle-conservation symmetry. We use the U(1) East, which lacks inversion symmetry, and the inversion-symmetric U(1) East-West models to illustrate our two main results. First, we observe that the simultaneous presence of constraints and chiral symmetry generally leads to a parametric increase in the number of zero modes due to the fragmentation of the many-body Hilbert space into disconnected sectors. Second, we generalize the concept of compact localized states from single-particle physics and introduce the notion of collective bound states, a special kind of nonergodic eigenstates that are robust to enlarging the system size. We formulate sufficient criteria for their existence, arguing that the degenerate zero mode subspace plays a central role, and demonstrate bound states in both example models and in a two-dimensional model, the U(1) North-East, and in the pairflip model, a system without particle conservation. Our results motivate a systematic study of bound states and their relation to ergodicity breaking, transport, and other properties of quantum kinetically constrained models. }, author = {Nicolau Jimenez, Eulalia and Ljubotina, Marko and Serbyn, Maksym}, issn = {2691-3399}, journal = {PRX Quantum}, publisher = {American Physical Society}, title = {{Fragmentation, zero modes, and collective bound states in constrained models}}, doi = {10.1103/sl79-1xgb}, volume = {7}, year = {2026}, } @article{21483, abstract = {Embryogenesis in the model plant Arabidopsis thaliana provides a framework for understanding how cell polarity and patterning coordinate with hormonal signalling to establish the plant body plan. Following fertilisation, the zygote divides asymmetrically to generate apical and basal lineages, establishing the apical–basal axis that defines future shoot and root poles. Genetic and molecular analyses of classical mutants including gnom, monopteros (mp), bodenlos (bdl) and topless revealed that localised auxin biosynthesis, directional transport and downstream transcriptional responses are central to apical–basal axis establishment and organ initiation. The main components of this regulation are polarly localised PIN auxin transporters and downstream modules involving MONOPTEROS and WUSCHEL-RELATED HOMEOBOX transcription factors. Advances in microscopy have transformed the study of Arabidopsis embryogenesis: fluorescence-compatible clearing reagents and three-dimensional reconstructions now permit quantitative analyses of cell geometry, division orientation, and cytoskeletal dynamics. Live ovule imaging setups with confocal laser scanning and multiphoton microscopes enable real-time observation of embryo development, while laser-assisted cell ablation can be used to probe cell-to-cell communication and fate plasticity. Together, these methodological breakthroughs position Arabidopsis embryos as a prime model for dissecting the chemical and biophysical cues that shape plant development.}, author = {Babic, David and Zupunski, Milan and Friml, Jiří}, issn = {1469-8137}, journal = {New Phytologist}, publisher = {Wiley}, title = {{Imaging and genetic toolbox to study Arabidopsis embryogenesis}}, doi = {10.1111/nph.71072}, year = {2026}, } @article{21502, abstract = {The mammalian brain stores glucose, the main circulating energy substrate, as glycogen. In rodents, the cerebellum contains relatively high glycogen levels, yet its cellular and subcellular distribution remains poorly defined. Using monoclonal antibodies against glycogen, we examined its distribution in the mouse cerebellar cortex. Glycogen was predominantly localized to Bergmann glia (BG) processes in the molecular layer and was also detected in Purkinje cells (PCs), the principal cerebellar neurons. To assess the functional significance of cerebellar glycogen, we analyzed behavior in mice lacking glycogen synthase 1 (Gys1) in BG or PCs using a floxed Gys1 line. Gys1 deficiency in either PCs or GFAP-positive cells reduced anxiety-like behavior, whereas combined deletion caused PC degeneration and ataxia. These findings reveal a critical role for glycogen metabolism in both astrocytes and neurons in cerebellar function.}, author = {Akther, Sonam and Lee, Ashley Bomin and Konno, Ayumu and Asiminas, Antonis and Vittani, Marta and Mishima, Tsuneko and Hirai, Hirokazu and Meehan, Claire Francesca and Duran, Jordi and Guinovart, Joan and Ashida, Hitoshi and Morita, Tsuyoshi and Baba, Otto and Shigemoto, Ryuichi and Nedergaard, Maiken and Hirase, Hajime}, issn = {2589-0042}, journal = {iScience}, number = {4}, publisher = {Elsevier}, title = {{Distribution and functional significance of rodent cerebellar glycogen}}, doi = {10.1016/j.isci.2026.115192}, volume = {29}, year = {2026}, } @phdthesis{20964, author = {Vladimirtsev, Dmitrii}, issn = {2791-4585}, pages = {22}, publisher = {Institute of Science and Technology Austria}, title = {{Armadillo repeat only proteins are master regulators of plant cyclic-nucleotide gated channels}}, doi = {10.15479/AT-ISTA-20964}, year = {2026}, } @phdthesis{21198, abstract = {In recent years there has been a massive increase in the amount of data generated in a decentralized manner. Ever more powerful edge devices, such as smartphones, have become ubiquitous in most societies on earth. Through text typed, photos taken and apps used, these devices, which we refer to as clients, generate enormous amounts of high quality and complex data. Moreover, the nature of these devices means the data they generate is often sensitive and privacy concerns prevent it being gathered and stored in a central location. This presents a challenge to the modern machine learning paradigm that requires central access to large amounts of data. Federated learning (FL) has emerged as one of the answers to this problem. Rather than bringing the data to the model, FL sends the model to the data. Model training takes place on device, with periodically synchronized updates, allowing data to remain locally stored. While this approach offers significant privacy advantages it comes with its own set of unique challenges. These include: data heterogeneity, the notion that different devices generate data in distinct ways which can negatively impact training dynamics; systems heterogeneity, meaning that different devices may have differing hardware specifications; high communication costs, which are induced by the repeated transferring of models over the network and low device computational power, which limits the use of larger models on device. In this thesis we present a range of methods for federated learning. We focus primarily on the challenge of data heterogeneity, though the methods presented are designed to be well adapted to the other challenges of a federated setting, such as the constraints of limited compute and communication overhead. We first present a method for explicitly modeling client data heterogeneity. The approach formulates clients as samples from a certain probability distribution and infers the parameters of this distribution from the available training clients. This learned distribution then represents the heterogeneity present among the clients and can be sampled from in order to create new simulated clients that are similar to the real clients we have observed so far. Following this we present two methods for directly dealing with data heterogeneity through personalization. Highly heterogeneous client data distributions can mean that learning a single global model becomes suboptimal, and some form of personalization of models to each individual client is required. Our approaches are based around hypernetworks, which we use to generate personalized model parameters without the need for additional training or finetuning. In the first approach we focus on generating full parameterizations of client models using learned embeddings of client data and labels, with a hypernetwork located on the central server. In the second approach we address the more challenging scenario where we want to generate a personalized model for a client without any label information. The hypernetwork is trained to generate a low dimensional representation of a client’s personalized model parameters, allowing it to be transferred to and run on the client devices. In our final presented method, we change our focus and rather than aim to directly address the challenge of data heterogeneity, we instead ensure we are unaffected by it. This is done in the context of k-means clustering and we present a method for federated clustering with a focus on added privacy guarantees.}, author = {Scott, Jonathan A}, issn = {2663-337X}, pages = {158}, publisher = {Institute of Science and Technology Austria}, title = {{Data heterogeneity and personalization in federated learning}}, doi = {10.15479/AT-ISTA-21198}, year = {2026}, } @phdthesis{21021, abstract = {This thesis examines how geometry and topology intersect in the representation, transformation, and analysis of complex shapes. It considers how continuous manifolds relate to their discrete analogues, how topological structures evolve in persistence vineyards, and how tools from topological data analysis can illuminate problems in mathematical physics. Central to this exploration is the question of how structure, both geometric and topological, persists or changes under approximation, sampling, or deformation. The work develops new approaches to skeletal and grid-based representations of surfaces, reveals the full expressive capacity of persistence vineyards, and applies topological methods to the longstanding problem of equilibria in electrostatic fields. These threads braid together into a broader understanding of how topology and geometry inform one another across theory, computation, and application.}, author = {Fillmore, Christopher D}, issn = {2663-337X}, pages = {122}, publisher = {Institute of Science and Technology Austria}, title = {{Braiding geometry and topology to study shapes and data}}, doi = {10.15479/AT-ISTA-21021}, year = {2026}, } @unpublished{21051, abstract = {In this work, we introduce and study what we believe is an intriguing and, to the best of our knowledge, previously unknown connection between two areas in computational topology, topological data analysis (TDA) and knot theory. Given a function from a topological space to $\mathbb{R}$, TDA provides tools to simplify and study the importance of topological features: in particular, the $l^{th}$-dimensional persistence diagram encodes the $l$-homology in the sublevel set as the function value increases as a set of points in the plane. Given a continuous one-parameter family of such functions, we can combine the persistence diagrams into an object known as a vineyard, which track the evolution of points in the persistence diagram. If we further restrict that family of functions to be periodic, we identify the two ends of the vineyard, yielding a closed vineyard. This allows the study of monodromy, which in this context means that following the family of functions for a period permutes the set of points in a non-trivial way. In this work, given a link and value $l$, we construct a topological space and periodic family of functions such that the closed $l$-vineyard contains this link. This shows that vineyards are topologically as rich as one could possibly hope. Importantly, it has at least two immediate consequences: First, monodromy of any periodicity can occur in a $l$-vineyard, answering a variant of a question by [Arya et al 2024]. To exhibit this, we also reformulate monodromy in a more geometric way, which may be of interest in itself. Second, distinguishing vineyards is likely to be difficult given the known difficulty of knot and link recognition, which have strong connections to many NP-hard problems.}, author = { Chambers, Erin and Fillmore, Christopher D and Stephenson, Elizabeth R and Wintraecken, Mathijs}, booktitle = {arXiv}, title = {{Braiding vineyards}}, doi = {10.48550/ARXIV.2504.11203}, year = {2026}, } @unpublished{21291, abstract = {The complexity and specificity of movement in vertebrates is driven by a rich diversity of spinal motor and interneuron cell types. During development, eleven spinal cord progenitor domains generate an equivalent number of cardinal neuron types. How progenitor domains, individual progenitors, and post-mitotic diversity relate is still unknown. We performed high-resolution, single-progenitor cell lineage tracing in the embryonic mouse spinal cord using mosaic analysis with double markers (MADM). Our quantitative study of lineage progression revealed that spinal cord progenitors undergo highly variable numbers of proliferative, neurogenic, and gliogenic cell divisions. The nascent clonally-related neurons migrate radially over large distances, span the dorsoventral axis, and even cross the midline, demonstrating striking bilaterality. Molecular and morphometric analysis indicate high levels of progenitor multipotency, with an individual progenitor capable of producing several molecularly and morphologically distinct neuron types, as well as astrocytes. These findings redefine spinal cord development as a process in which lineage variability — rather than rigid progenitor identity — drives the generation of cellular diversity.}, author = {Gobeil, Sophie A and Da Silveira Neto, Francisco and Silvestrelli, Giulia and Smits, Matthijs Geert and Streicher, Carmen and Cheung, Giselle T and Hippenmeyer, Simon and Sweeney, Lora Beatrice Jaeger}, booktitle = {bioRxiv}, title = {{Lineage origin of spinal cord cell type diversity}}, doi = {10.64898/2026.02.12.705305}, year = {2026}, } @inproceedings{21134, abstract = {The Nakamoto consensus protocol underlying the Bitcoin blockchain uses proof of work as a voting mechanism. Honest miners who contribute hashing power towards securing the chain try to extend the longest chain they are aware of. Despite its simplicity, Nakamoto consensus achieves meaningful security guarantees assuming that at any point in time, a majority of the hashing power is controlled by honest parties. This also holds under “resource variability”, i.e., if the total hashing power varies greatly over time. Proofs of space (PoSpace) have been suggested as a more sustainable replacement for proofs of work. Unfortunately, no construction of a “longest-chain” blockchain based on PoSpace, that is secure under dynamic availability, is known. In this work, we prove that without additional assumptions no such protocol exists. We exactly quantify this impossibility result by proving a bound on the length of the fork required for double spending as a function of the adversarial capabilities. This bound holds for any chain selection rule, and we also show a chain selection rule (albeit a very strange one) that almost matches this bound. The Nakamoto consensus protocol underlying the Bitcoin blockchain uses proof of work as a voting mechanism. Honest miners who contribute hashing power towards securing the chain try to extend the longest chain they are aware of. Despite its simplicity, Nakamoto consensus achieves meaningful security guarantees assuming that at any point in time, a majority of the hashing power is controlled by honest parties. This also holds under “resource variability”, i.e., if the total hashing power varies greatly over time. Proofs of space (PoSpace) have been suggested as a more sustainable replacement for proofs of work. Unfortunately, no construction of a “longest-chain” blockchain based on PoSpace, that is secure under dynamic availability, is known. In this work, we prove that without additional assumptions no such protocol exists. We exactly quantify this impossibility result by proving a bound on the length of the fork required for double spending as a function of the adversarial capabilities. This bound holds for any chain selection rule, and we also show a chain selection rule (albeit a very strange one) that almost matches this bound. Concretely, we consider a security game in which the honest parties at any point control 0 > 1 times more space than the adversary. The adversary can change the honest space by a factor 1+- E with every block (dynamic availability), and “replotting” the space (which allows answering two challenges using the same space) takes as much time as p blocks. We prove that no matter what chain selection rule is used, in this game the adversary can create a fork of length o^2 . p/E that will be picked as the winner by the chain selection rule. We also provide an upper bound that matches the lower bound up to a factor o. There exists a chain selection rule (albeit a very strange one) which in the above game requires forks of length at least o . p/E Our results show the necessity of additional assumptions to create a secure PoSpace based longest-chain blockchain. The Chia network in addition to PoSpace uses a verifiable delay function. Our bounds show that an additional primitive like that is necessary.}, author = {Baig, Mirza Ahad and Pietrzak, Krzysztof Z}, booktitle = {29th International Conference on Financial Cryptography and Data Security}, isbn = {9783032070340}, issn = {1611-3349}, location = {Miyakojima, Japan}, pages = {127--142}, publisher = {Springer Nature}, title = {{On the (in)security of Proofs-of-space based longest-chain blockchains}}, doi = {10.1007/978-3-032-07035-7_8}, volume = {15752}, year = {2026}, } @phdthesis{21651, abstract = {Blockchains enable distributed consensus in permissionless settings, where participants are unknown, dynamically changing, and do not trust each other. While Bitcoin, based on Proof-of-Work (PoW), was the first protocol in this model, significant research has focused on permissionless protocols using alternative physical resources, specifically Proof-of-Space (PoSpace) and Verifiable Delay Functions (VDFs). This thesis investigates the theoretical limits and design space of longest-chain protocols in the fully permissionless and dynamically available settings using these three resources. First, we address the feasibility of blockchains relying solely on storage as a resource. We prove a fundamental impossibility result: there exists no secure longest-chain protocol based exclusively on Proof-of-Space in the fully permissionless or dynamically available settings. Further, we quantify the adversarial capabilities required to execute a double-spend attack. Our result formally justifies the necessity of coupling PoSpace with time-dependent primitives (such as VDFs) or to move to less permissive settings (quasi-permissionless or permissioned) to ensure security. Second, we generalize Nakamoto-like heaviest chain consensus to protocols utilizing combinations of multiple physical resources. We analyze chain selection rules governed by a weight function Γ(S, V,W), which assigns weight to blocks based on recorded Space (S), VDF speed (V ), and Work (W). We provide a complete classification of secure weight functions, proving that a weight function is secure against private double-spend attacks if and only if it is homogeneous in the timed resources (V,W) and sub-homogeneous in S. This framework unifies existing protocols like Bitcoin and Chia under a single theoretical model and provides a powerful tool for designing new longest-chain blockchains from a mix of physical resources.}, author = {Baig, Mirza Ahad}, isbn = {978-3-99078-078-7}, issn = {2663-337X}, publisher = {Institute of Science and Technology Austria}, title = {{On secure chain selection rules from physical resources in a permissionless setting}}, doi = {10.15479/AT-ISTA-21651}, year = {2026}, } @phdthesis{20991, abstract = {Rapid local adaptation to new environments is critical for species persistence, especially in introduced populations. The evolutionary success of these populations is fundamentally dictated by the organization of genetic variation—the genomic architecture—in the face of severe demographic constraints, such as the founder effects and genetic bottlenecks that frequently accompany colonization. A central question in evolutionary biology is whether rapid adaptation relies on major-effect loci, such as chromosomal inversions, or on many small-effect loci dispersed across the genome. Furthermore, the genomic architecture strongly influences the extent to which evolutionary outcomes are predictable. Using introduced populations of the marine snail, Littorina saxatilis, as a model, this thesis investigates how genetic variation and genomic structure drive adaptation following introduction. We employed a population genomics approach on experimentally and accidentally introduced populations to dissect the specific genomic features that underpin divergence in newly colonized environments. In Chapter 2, we tested the predictability of local adaptation through an uncommon 30-year transplant experiment in nature. By distinguishing allele and chromosomal inversion frequency changes from neutral expectations, we found that evolutionary change was highly predictable at the macro-scale (phenotypes and chromosomal inversions), but less robust at the level of individual collinear loci. This result demonstrates that evolution can be predictable when a population possesses sufficient standing genetic variation (SGV), with chromosomal inversions acting as key integrated units that facilitate a rapid response to selection. Building on this, Chapter 3 applied whole-genome sequencing to three accidentally introduced populations (Venice, San Francisco, and Redwood City) to investigate their likely source and genomic patterns of divergence. We identified genomic regions of remarkable divergence potentially associated with local adaptation, and likely fuelled by SGV, while explicitly acknowledging the difficulty in disentangling selection signals from the genome-wide effects of demographic processes. Furthermore, we found that the divergence patterns relied extensively on the collinear genome in these introduced populations, and less clearly on the chromosomal inversions. This observation contrasts with local adaptation observed in the experimental system that relied on both collinear loci and highly selected chromosomal inversions, highlighting how demographic history and genomic architecture influence the detectable signature of local adaptation. A major limitation to conducting large-scale comparative evolutionary studies is the lack of data standardization, which prevents the integration of community knowledge and high-resolution environmental and genetic data. Chapter 4 addresses this by developing a community database for the Littorina system. This platform implements standardized protocols for the integration of diverse phenotypic and environmental data from multiple Littorina species. Likewise, the platform also centralizes the availability of associated genomic data through links to external repositories. This database represents a crucial tool to test complex, large-scale evolutionary hypotheses. Collectively, this thesis strongly reinforces the fundamental importance of SGV as the raw material for successful local adaptation, a conclusion supported by evidence in both experimental and accidental introductions. Furthermore, this work highlights the critical role of the genomic architecture—specifically chromosomal inversions—in driving the predictability and effectiveness of adaptive responses. Our findings underscore how the interplay between SGV and genomic architecture dictates the trajectory and detectability of evolution in colonizing populations, while simultaneously providing a necessary tool to advance comparative evolutionary genomics in emerging model organisms.}, author = {Garcia Castillo, Diego Fernando}, isbn = {978-3-99078-077-0}, issn = {2663-337X}, pages = {199}, publisher = {Institute of Science and Technology Austria}, title = {{The genomic architecture of local adaptation in introduced populations}}, doi = {10.15479/AT-ISTA-20991}, year = {2026}, } @article{21765, abstract = {Dielectric particles of the same material exchange electrical charge during collisions or sliding contacts, yet the underlying charge-exchange mechanism is still not understood. The fact that particles can become highly charged as a result of this effect has significant consequences for many settings, both in nature and industry, such as thunderstorms, volcanic eruptions, particle aggregation during meteorite and planet formation, and the clogging of industrial granular systems. Toward understanding these systems, great efforts have been made to develop precise in situ measurements for particle charge, e.g., to determine ensemble charge distributions or measure exchange during individual contacts. Here, we present experimental results concerning the particle size scaling of the stationary-state charge distributions of oxide particles in the sub-millimeter range. We measure the charge distributions for large ensembles of monodisperse ZrO2:SiO2 composite spheres, ranging from 172 to 545µ⁢m in diameter. These distributions are non-Gaussian and collapse to a single master curve when plotted as functions of the surface charge density Σ=𝑞/4⁢𝜋⁢𝑅2. X-ray fluorescence and atomic force microscopy measurements show that the differences in the measured charge distributions are not due to variations in chemical composition or surface roughness, but rather to size alone. Our findings provide constraints on microscopic models for charge exchange, namely that they should lead to steady-state distributions that are non-Gaussian and scale in a specific way with particle size.}, author = {Lara, Macarena and Flores, Marcos and Castillo, Gustavo and Tassara, Santiago and Waitukaitis, Scott R and Mujica, Nicolás}, issn = {2475-9953}, journal = {Physical Review Materials}, number = {4}, publisher = {American Physical Society}, title = {{Particle size scaling of non-Gaussian granular charge distributions}}, doi = {10.1103/qw6t-xqdw}, volume = {10}, year = {2026}, } @article{21485, abstract = {Insulating oxides are among the most abundant solid materials in the universe1,2,3. Of the many ways in which they influence natural phenomena, perhaps the most consequential is their capacity to transfer electrical charge during contact4,5,6,7,8,9,10—which occurs even between samples of the same oxide—yet the symmetry-breaking parameter that causes this remains unidentified11,12. Here we show that adventitious carbonaceous molecules adsorbed from the environment are the symmetry-breaking factor in same-material oxide contact electrification (CE). We use acoustic levitation to measure charge exchange between a sphere and a plate composed of identical amorphous silicon dioxide (SiO2). Although charging polarity is random for co-prepared samples, we control it with baking or plasma treatment. Observing the charge-exchange relaxation afterwards, we see dynamics over a timescale of hours and connect this directly to the presence of adventitious carbon with time-of-flight mass spectrometry, low-energy ion scattering and infrared spectroscopy. Going further, we confirm that adventitious carbon can even determine charge exchange among different oxides. Our results identify the symmetry-breaking parameter that causes insulating oxides to exchange charge in settings ranging from desert sands4 to volcanic plumes5,6, while simultaneously highlighting an overlooked factor in CE more broadly.}, author = {Grosjean, Galien M and Ostermann, Markus and Sauer, Markus and Hahn, Michael and Pichler, Christian M. and Fahrnberger, Florian and Pertl, Felix and Balazs, Daniel and Link, Mason M. and Kim, Seong H. and Schrader, Devin L. and Blanco, Adriana and Gracia, Francisco and Mujica, Nicolás and Waitukaitis, Scott R}, issn = {1476-4687}, journal = {Nature}, number = {8106}, pages = {626--631}, publisher = {Springer Nature}, title = {{Adventitious carbon breaks symmetry in oxide contact electrification}}, doi = {10.1038/s41586-025-10088-w}, volume = {651}, year = {2026}, } @article{21015, abstract = {Early embryo geometry is one of the most invariant species-specific traits, yet its role in ensuring developmental reproducibility and robustness remains underexplored. Here we show that in zebrafish, the geometry of the fertilized egg—specifically its curvature and volume—serves as a critical initial condition triggering a cascade of events that influence development. The embryo geometry guides patterned asymmetric cell divisions in the blastoderm, generating radial gradients of cell volume and nucleocytoplasmic ratio. These gradients generate mitotic phase waves, with the nucleocytoplasmic ratio determining individual cell cycle periods independently of other cells. We demonstrate that reducing cell autonomy reshapes these waves, emphasizing the instructive role of geometry-derived volume patterns in setting the intrinsic period of the cell cycle oscillator. In addition to organizing cell cycles, early embryo geometry spatially patterns zygotic genome activation at the midblastula transition, a key step in establishing embryonic autonomy. Disrupting the embryo shape alters the zygotic genome activation pattern and causes ectopic germ layer specification, underscoring the developmental significance of geometry. Together, our findings reveal a symmetry-breaking function of early embryo geometry in coordinating cell cycle and transcriptional patterning.}, author = {Mishra, Nikhil and Li, Yuting I and Hannezo, Edouard B and Heisenberg, Carl-Philipp J}, issn = {1745-2481}, journal = {Nature Physics}, pages = {139--150}, publisher = {Springer Nature}, title = {{Geometry-driven asymmetric cell divisions pattern cell cycles and zygotic genome activation in the zebrafish embryo}}, doi = {10.1038/s41567-025-03122-1}, volume = {22}, year = {2026}, } @article{21382, abstract = {The exceptional energy-harvesting efficiency of lead-halide perovskites arises from unusually long photocarrier diffusion lengths and recombination lifetimes that persist even in defect-rich, solution-grown samples. Paradoxically, perovskites are also known for having very short exciton decay times. Here, we resolve this apparent contradiction by showing that key optoelectronic properties of perovskites can be explained by localized flexoelectric polarization confined to interfaces between domains of spontaneous strain. Using birefringence imaging, electrochemical staining, and zero-bias photocurrent measurements, we visualize the domain structure and directly probe the associated internal fields in nominally cubic single crystals of methylammonium lead bromide. We demonstrate that localized flexoelectric fields spatially separate electrons and holes to opposite sides of domain walls, exponentially suppressing recombination. Domain walls thus act as efficient mesoscopic transport channels for long-lived photocarriers, microscopically linking structural heterogeneity to charge transport and offering mechanistically informed design principles for perovskite solar-energy technologies.}, author = {Rak, Dmytro and Lorenc, Dusan and Balazs, Daniel and Zhumekenov, Ayan A. and Bakr, Osman M. and Alpichshev, Zhanybek}, issn = {2041-1723}, journal = {Nature Communications}, publisher = {Springer Nature}, title = {{Flexoelectric domain walls enable charge separation and transport in cubic perovskites}}, doi = {10.1038/s41467-026-68660-5}, volume = {17}, year = {2026}, } @article{21759, abstract = {Promoters and enhancers are cis-regulatory elements (CREs), DNA sequences that bind transcription factor (TF) proteins to up- or down-regulate target genes. Decades-long efforts yielded TF-DNA interaction models that predict how strongly an individual TF binds arbitrary DNA sequences and how individual binding events on the CRE combine to affect gene expression. These insights can be synthesized into a global, biophysically realistic, and quantitative genotype-phenotype (GP) map for gene regulation, a ‘holy grail’ for the application of evolutionary theory. A global map provides a rare opportunity to simulate the long-term evolution of regulatory sequences and pose several fundamental questions: How long does it take to evolve CREs de novo? How many non-trivial regulatory functions exist in sequence space? How connected are they? For which regulatory architecture is CRE evolution most rapid and evolvable? In this article, the second of a two-part series, we review the application of evolutionary concepts — epistasis, robustness, evolvability, tunability, plasticity, and bet-hedging — to the evolution of gene regulatory sequences. We then evaluate the potential for a unifying theory for the evolution of regulatory sequences and identify key open challenges.}, author = {Mascolo, Elia and Körei, Reka E and Borst, Noa O. and Barton, Nicholas H and Crocker, Justin and Tkačik, Gašper}, issn = {1879-0380}, journal = {Current Opinion in Genetics and Development}, publisher = {Elsevier}, title = {{Long-term evolution of regulatory DNA sequences. Part 2: Theory and future challenges}}, doi = {10.1016/j.gde.2026.102472}, volume = {98}, year = {2026}, } @article{21762, abstract = {Bacteria, like eukaryotes, use conserved cytoskeletal systems for intracellular organization. The plasmid-encoded ParMRC system forms actin-like filaments that segregate low–copy number plasmids. In multicellular cyanobacteria such as Anabaena sp., we found that a chromosomally encoded ParMR system has evolved into a cytoskeletal system named CorMR with a function in cell shape control rather than DNA segregation. Live-cell imaging, in vitro reconstitution, and cryo–electron microscopy revealed that CorM formed dynamically unstable, antiparallel double-stranded filaments that were recruited to the membrane by CorR through an amphipathic helix conserved in multicellular cyanobacteria. CorMR filaments were regulated by MinC, which excluded them from the poles and division plane. Comparative genomics indicated that the repurposing of ParMR and Min systems coevolved with cyanobacterial multicellularity, highlighting the evolutionary plasticity of cytoskeletal systems in bacteria.}, author = {Springstein, Benjamin L and Javoor, Manjunath and Megrian, Daniela and Hajdu, Roman and Hanke, Dustin M. and Zens, Bettina and Weiss, Gregor L. and Schur, Florian Km and Loose, Martin}, issn = {1095-9203}, journal = {Science}, number = {6795}, publisher = {AAAS}, title = {{Repurposing of a DNA segregation machinery into a cytoskeletal system controlling cell shape}}, doi = {10.1126/science.aea6343}, volume = {392}, year = {2026}, } @article{21704, abstract = {How functional protein sequences are distributed in sequence space is fundamentally important for evolutionary theory and protein design, particularly if a large diversity of protein functions are hidden in evolutionarily unexplored areas of the sequence space. However, this question is understudied in part because experimental and computational studies use extant sequences as a starting point to study sequence space. Here, we study whether extant sequences are representative of the entire functional sequence space. Across thousands of protein families from vertebrates and bacteria we calculate the dimensionality and the volume of sequence space occupied by extant homologs. We find that the observed dimensionality and volume of extant sequence space are minuscule, many orders of magnitude smaller than what we estimated using a model of protein evolution. Simulating sequence evolution we then quantify the impact of phylogeny, selection, and epistasis on restricting the evolutionary exploration of sequence space. We find that sequence evolution from a single common ancestor, or a single point of origin in sequence space, is by far the largest limiting factor that reduces the dimensionality and volume of extant sequence space. These results indicate that there are vast areas of functional sequence space that have not been explored in evolution because of the excessive restrictions on natural exploration of the protein sequence space imposed by the point of origin effect. We suggest that protein design methods that rely on extant sequences may be limited in their ability to discover truly novel functions.}, author = {Isakova, Lada H. and Streltsova, Elizaveta and Bochkareva, Olga and Vlasov, Peter K. and Kondrashov, Fyodor}, issn = {1091-6490}, journal = {Proceedings of the National Academy of Sciences}, number = {14}, pages = {e2532018123}, publisher = {National Academy of Sciences}, title = {{Descent from a common ancestor restricts exploration of protein sequence space}}, doi = {10.1073/pnas.2532018123}, volume = {123}, year = {2026}, } @article{21746, abstract = {As vertebrates transitioned from water to land, locomotion shifted from undulatory swimming to limb-based movement. How spinal circuits and their cell types evolved to support this transition remains unclear. We leverage frog metamorphosis, which recapitulates this transition within a single organism, to define how spinal circuits generate aquatic versus terrestrial motor patterns. At swim stages, spinal architecture is uniform, with a transcriptionally and anatomically homogeneous motor and interneurons. As limbs develop and their movement complexifies, spinal circuits expand in neuron number and subtype diversity. This expansion is most pronounced for V1 inhibitory neurons, which increase ∼70-fold and diversify into transcriptionally distinct subtypes. Disrupting transcription factors defining emerging motor and V1 populations reveals molecular segregation between swim and limb circuits, highlighting the role of subtype diversity in motor coordination. A multifold increase in inhibitory neuron diversity thus underlies the tail-to-limb locomotor transition, providing a framework for spinal circuit adaptation during vertebrate evolution.}, author = {Vijatovic, David and Toma, Florina Alexandra and Ignatyev, Y and Harrington, Zoe P and Sommer, Christoph M and Hauschild, Robert and Smits, Matthijs Geert and Dalla Vecchia, Marco and Trevisan, Alexandra J. and Chapman, Phillip and Julseth, Mara and Brenner-Morton, Susan and Gabitto, Mariano I. and Dasen, Jeremy S. and Bikoff, Jay B. and Sweeney, Lora Beatrice Jaeger}, issn = {2211-1247}, journal = {Cell Reports}, number = {4}, publisher = {Elsevier}, title = {{Multifold increase in spinal inhibitory cell types with emergence of limb movement}}, doi = {10.1016/j.celrep.2026.117227}, volume = {45}, year = {2026}, }