Irep Gözen

• Katke, Chinmay; Pedrueza-Villalmanzo, Esteban; Spustova, Karolina; Ryskulov, Ruslan; Kaplan, C. Nadir & Gözen, Irep (2023). Colony-like Protocell Superstructures. ACS Nano. ISSN 1936-0851. 17(4), p. 3368–3382. doi: 10.1021/acsnano.2c08093. Full text in Research Archive
• Xue, Lin; Stephenson, Anna B. & Gözen, Irep (2022). Liposome-assisted in-situ cargo delivery to artificial cells and cellular subcompartments. bioRxiv. ISSN 2692-8205. doi: 10.1101/2022.04.27.489538.
• Schanke, Ingrid Jin; Xue, Lin; Spustova, Karolina & Gözen, Irep (2022). Transport among protocells via tunneling nanotubes. Nanoscale. ISSN 2040-3364. 14(29), p. 10418–10427. doi: 10.1039/d2nr02290g. Full text in Research Archive
• Gözen, Irep; Köksal, Elif; Põldsalu, Inga; Xue, Lin; Spustova, Karolina & Pedrueza-Villalmanzo, Esteban [Show all 9 contributors for this article] (2022). Protocells: Milestones and Recent Advances. Small. ISSN 1613-6810. 18(18). doi: 10.1002/smll.202106624. Full text in Research Archive Show summary

  The origin of life is still one of humankind's great mysteries. At the transition between nonliving and living matter, protocells, initially featureless aggregates of abiotic matter, gain the structure and functions necessary to fulfill the criteria of life. Research addressing protocells as a central element in this transition is diverse and increasingly interdisciplinary. The authors review current protocell concepts and research directions, address milestones, challenges and existing hypotheses in the context of conditions on the early Earth, and provide a concise overview of current protocell research methods.

• Köksal, Elif; Põldsalu, Inga; Friis, Henrik; Mojzsis, Stephen J; Bizzarro, Martin & Gözen, Irep (2022). Spontaneous formation of prebiotic compartment colonies on Hadean Earth and pre-Noachian Mars. ChemSystemsChem. ISSN 2570-4206. 4(3). doi: 10.1002/syst.202100040. Full text in Research Archive
• Spustova, Karolina; Xue, Lin; Ryskulov, Ruslan; Jesorka, Aldo & Gözen, Irep (2021). Manipulation of Lipid Membranes with Thermal Stimuli, Membrane Lipids. Springer. ISSN 978-1-0716-1842-4. p. 209–225.
• Põldsalu, Inga; Köksal, Elif & Gözen, Irep (2021). Mixed fatty acid-phospholipid protocell networks. bioRxiv. ISSN 2692-8205. doi: 10.1101/2021.03.08.434432.
• Gözen, Irep (2021). Did Solid Surfaces Enable the Origin of Life? Life. ISSN 2075-1729. doi: 10.3390/life11080795. Full text in Research Archive
• Schanke, Ingrid Jin; Xue, Lin; Spustova, Karolina & Gözen, Irep (2021). Transport among protocells via tunneling nanotubes. bioRxiv. ISSN 2692-8205. doi: 10.1101/2021.09.16.460285.
• Spustova, Karolina; Katke, Chinmay; Villalmanzo, Esteban Pedrueza; Ryskulov, Ruslan; Kaplan, C. Nadir & Gözen, Irep (2021). Colony-like Protocell Superstructures. bioRxiv. ISSN 2692-8205. doi: 10.1101/2021.09.16.460583.
• Põldsalu, Inga; Köksal, Elif & Gözen, Irep (2021). Mixed fatty acid-phospholipid protocell networks. Physical Chemistry, Chemical Physics - PCCP. ISSN 1463-9076. 23(47), p. 26948–26954. doi: 10.1039/d1cp03832j. Full text in Research Archive
• Köksal, Elif; Liese, Susanne; Xue, Lin; Ryskulov, Ruslan; Viitala, Lauri & Carlson, Andreas [Show all 7 contributors for this article] (2020). Rapid Growth and Fusion of Protocells in Surface‐Adhered Membrane Networks. Small. ISSN 1613-6810. 16(38). doi: 10.1002/smll.202002529. Show summary

  Elevated temperatures might have promoted the nucleation, growth, and replication of protocells on the early Earth. Recent reports have shown evidence that moderately high temperatures not only permit protocell assembly at the origin of life, but can have actively supported it. Here, the fast nucleation and growth of vesicular compartments from autonomously formed lipid networks on solid surfaces, induced by a moderate increase in temperature, are shown. Branches of the networks, initially consisting of self‐assembled interconnected nanotubes, rapidly swell into microcompartments which can spontaneously encapsulate RNA fragments. The increase in temperature further causes fusion of adjacent network‐connected compartments, resulting in the redistribution of the RNA. The experimental observations and the mathematical model indicate that the presence of nanotubular interconnections between protocells facilitates the fusion process.

• Gözen, Irep & Dommersnes, Paul Gunnar (2020). Biological lipid nanotubes and their potential role in evolution. The European Physical Journal Special Topics. ISSN 1951-6355. 229(17-18), p. 2843–2862. doi: 10.1140/epjst/e2020-000130-7. Full text in Research Archive Show summary

  The membrane of cells and organelles are highly deformable fluid interfaces, and can take on a multitude of shapes. One distinctive and particularly interesting property of biological membranes is their ability to from long and uniform nanotubes. These nanoconduits are surprisingly omnipresent in all domains of life, from archaea, bacteria, to plants and mammals. Some of these tubes have been known for a century, while others were only recently discovered. Their designations are different in different branches of biology, e.g. they are called stromule in plants and tunneling nanotubes in mammals. The mechanical transformation of flat membranes to tubes involves typically a combination of membrane anchoring and external forces, leading to a pulling action that results in very rapid membrane nanotube formation – micrometer long tubes can form in a matter of seconds. Their radius is set by a mechanical balance of tension and bending forces. There also exists a large class of membrane nanotubes that form due to curvature inducing molecules. It seems plausible that nanotube formation and functionality in plants and animals may have been inherited from their bacterial ancestors during endosymbiotic evolution. Here we attempt to connect observations of nanotubes in different branches of biology, and outline their similarities and differences with the aim of providing a perspective on their joint functions and evolutionary origin.

• Spustová, Karolina; Köksal, Elif; Ainla, Alar & Gözen, Irep (2020). Subcompartmentalization and Pseudo‐Division of Model Protocells. Small. ISSN 1613-6810. 17(2). doi: 10.1002/smll.202005320. Show summary

  Membrane enclosed intracellular compartments have been exclusively associated with the eukaryotes, represented by the highly compartmentalized last eukaryotic common ancestor. Recent evidence showing the presence of membranous compartments with specific functions in archaea and bacteria makes it conceivable that the last universal common ancestor and its hypothetical precursor, the protocell, may have exhibited compartmentalization. To the authors’ knowledge, there are no experimental studies yet that have tested this hypothesis. They report on an autonomous subcompartmentalization mechanism for protocells which results in the transformation of initial subcompartments to daughter protocells. The process is solely determined by the fundamental materials properties and interfacial events, and does not require biological machinery or chemical energy supply. In the light of the authors’ findings, it is proposed that similar events may have taken place under early Earth conditions, leading to the development of compartmentalized cells and potentially, primitive division.

• Joemetsa, Silver; Spustová, Karolina; Kustanovich, Kiryl; Ainla, Alar; Schindler, Severin & Eigler, Siegfried [Show all 10 contributors for this article] (2019). Molecular Lipid Films on Microengineering Materials. Langmuir. ISSN 0743-7463. 35(32), p. 10286–10298. doi: 10.1021/acs.langmuir.9b01120.
• Kustanovich, Kiryl; Yantchev, Ventsislav; Doosti, Baharan Ali; Gözen, Irep & Jesorka, Aldo (2019). A microfluidics-integrated impedance/surface acoustic resonance tandem sensor. Sensing and Bio-Sensing Research. 25. doi: 10.1016/j.sbsr.2019.100291.
• Rydmark, Marcella Orwick; Christensen, Mikkel; Köksal, Elif; Kantarci, Ilayda; Kustanovich, Kiryl & Yantchev, Ventsislav [Show all 8 contributors for this article] (2019). Styrene maleic acid copolymer induces pores in biomembranes . Soft Matter. ISSN 1744-683X. 15(39), p. 7934–7944. doi: 10.1039/C9SM01407A. Full text in Research Archive
• Gözen, Irep (2019). A Hypothesis for Protocell Division on the Early Earth. ACS Nano. ISSN 1936-0851. 13(10), p. 10869–10871. doi: 10.1021/acsnano.9b06584.
• Köksal, Elif; Belletati, Patrícia Fernandes; Reint, Ganna; Olsson, Ragni; Leitl, Kira & Kantarci, Ilayda [Show all 7 contributors for this article] (2019). Spontaneous Formation and Rearrangement of Artificial Lipid Nanotube Networks as a Bottom-Up Model for Endoplasmic Reticulum. Journal of Visualized Experiments. ISSN 1940-087X. 2019(143), p. 1–9. doi: 10.3791/58923. Full text in Research Archive
• Köksal, Elif; Liese, Susanne; Kantarci, Ilayda; Olsson, Ragni; Carlson, Andreas & Gözen, Irep (2019). Nanotube-Mediated Path to Protocell Formation. ACS Nano. ISSN 1936-0851. 13(6), p. 6867–6878. doi: 10.1021/acsnano.9b01646. Full text in Research Archive
• Horowitz, Viva R; Chambers, Zachary C; Gözen, Irep; Dimiduk, Thomas G & Manoharan, Vinothan N (2019). Active colloidal particles in emulsion droplets: a model system for the cytoplasm. The European Physical Journal Special Topics. ISSN 1951-6355. 227(17), p. 2413–2424. doi: 10.1140/epjst/e2019-800026-y. Show summary

  In living cells, molecular motors create activity that enhances the diffusion of particles throughout the cytoplasm, and not just ones attached to the motors. We demonstrate initial steps toward creating artificial cells that mimic this phenomenon. Our system consists of active, Pt-coated Janus particles and passive tracers confined to emulsion droplets. We track the motion of both the active particles and passive tracers in a hydrogen peroxide solution, which serves as the fuel to drive the motion. We first show that correcting for bulk translational and rotational motion of the droplets induced by bubble formation is necessary to accurately track the particles. After drift correction, we find that the active particles show enhanced diffusion in the interior of the droplets and are not captured by the droplet interface. At the particle and hydrogen peroxide concentrations we use, we observe little coupling between the active and passive particles. We discuss the possible reasons for lack of coupling and describe ways to improve the system to more effectively mimic cytoplasmic activity.

• Gupta, Abhay; Gözen, Irep & Taylor, Michael (2019). A cellular automaton for modeling non-trivial biomembrane ruptures. Soft Matter. ISSN 1744-683X. 15(20), p. 4178–4186. doi: 10.1039/c8sm02032a. Full text in Research Archive
• Bilal, Taylan & Gözen, Irep (2017). Formation and dynamics of endoplasmic reticulum-like lipid nanotube networks. Biomaterials Science. ISSN 2047-4830. 5(7), p. 1256–1264. doi: 10.1039/c7bm00227k.
• Taylor, Michael; Gözen, Irep; Patel, Samir; Jesorka, Aldo & Bertoldi, Katia (2016). Peridynamic Modeling of Ruptures in Biomembranes. PLOS ONE. ISSN 1932-6203. doi: 10.1371/journal.pone.0165947.
• Gözen, Irep & Jesorka, Aldo (2015). Lipid nanotube networks: Biomimetic Cell-to-Cell Communication and Soft-Matter Technology. Nanofabrication. ISSN 2299-680X. 2, p. 27–31. doi: 10.1515/nanofab-2015-0003.
• Luo, ZhengYuan; Güven, Sinan; Gözen, Irep; Chen, Pu; Tasoglu, Savas & Anchan, Raymond M [Show all 8 contributors for this article] (2015). Deformation of a single mouse oocyte in a constricted microfluidic channel. Microfluidics and Nanofluidics. ISSN 1613-4982. 19, p. 883–890. doi: 10.1007/s10404-015-1614-0.
• Gözen, Irep & Dommersnes, Paul Gunnar (2014). Pore dynamics in lipid membranes. The European Physical Journal Special Topics. ISSN 1951-6355. 223, p. 1813–1829. doi: 10.1140/epjst/e2014-02228-5.
• El Assal, Rami; Güven, Sinan; Gürkan, Umut Atakan; Gözen, Irep; Shafiee, Hadi & Dalbeyler, Sedef [Show all 18 contributors for this article] (2014). Bio-Inspired Cryo-Ink Preserves Red Blood Cell Phenotype and Function During Nanoliter Vitrification. Advanced Materials. ISSN 0935-9648. 26, p. 5815–5822. doi: 10.1002/adma.201400941.
• Gözen, Irep; Örtmen, Bahanur; Põldsalu, Inga; Dommersnes, Paul Gunnar; Orwar, Owe & Jesorka, Aldo (2013). Repair of large area pores in supported double bilayers. Soft Matter. ISSN 1744-683X. 9, p. 2787 –2792. doi: 10.1039/C3SM27429B.
• Ainla, Alar; Gözen, Irep; Hakonen, Bodil & Jesorka, Aldo (2013). Lab on a Biomembrane: Rapid prototyping and manipulation of 2D fluidic lipid bilayer circuits. Scientific Reports. ISSN 2045-2322. 3. doi: 10.1038/srep02743.
• Gözen, Irep; Shaali, Mehrnaz; Ainla, Alar; Örtmen, Bahanur; Põldsalu, Inga & Kustanovich, Kiryl [Show all 10 contributors for this article] (2013). Thermal migration of molecular lipid films as a contactless fabrication strategy for lipid nanotube networks. Lab on a Chip. ISSN 1473-0197. 13, p. 3822 –3826. doi: 10.1039/C3LC50391G.
• Gözen, Irep & Jesorka, Aldo (2012). Instrumental Methods to Characterize Molecular Phospholipid Films on Solid Supports. Analytical Chemistry. ISSN 0003-2700. 84 (2), p. 822–838. doi: 10.1021/ac203126f.
• Gözen, Irep; Dommersnes, Paul Gunnar; Orwar, Owe & Jesorka, Aldo (2012). Evidence for membrane flow through pores in stacked phospholipid membranes. Soft Matter. ISSN 1744-683X. 8, p. 6220 –6225. doi: 10.1039/C2SM25629K.
• Gözen, Irep; Billerit, Celine; Dommersnes, Paul Gunnar; Jesorka, Aldo & Orwar, Owe (2011). Calcium-ion-controlled nanoparticle-induced tubulation in supported flat phospholipid vesicles. Soft Matter. ISSN 1744-683X. 7, p. 9706–9713 . doi: 10.1039/C1SM05677H.
• Gözen, Irep & Örmeci, Banu (2010). Effect of daylight on regrowth of bacteria in anaerobically digested sludge. Water Science and Technology. ISSN 0273-1223. 62 (2) , p. 364–369. doi: 10.2166/wst.2010.268.
• Lobovkina, Tatsiana; Gözen, Irep; Erkan, Yavuz; Olofsson, Jessica; Weber, Stephen G. & Orwar, Owe (2010). Protrusive growth and periodic contractile motion in surface-adhered vesicles induced by Ca2+-gradients. Soft Matter. ISSN 1744-683X. 6, p. 268 –272. doi: 10.1039/B916805M.
• Gözen, Irep; Dommersnes, Paul Gunnar; Czolkos, Ilja; Jesorka, Aldo; Lobovkina, Tatsiana & Orwar, Owe (2010). Fractal avalanche ruptures in biological membranes. Nature Materials. ISSN 1476-1122. 9, p. 908–912. doi: 10.1038/nmat2854.
• Ainla, Alar; Gözen, Irep; Orwar, Owe & Jesorka, Aldo (2009). A Microfluidic Diluter Based on Pulse Width Flow Modulation. Analytical Chemistry. ISSN 0003-2700. 81 (13), p. 5549–5556. doi: 10.1021/ac9010028.

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• Gözen, Irep; Dommersnes, Paul Gunnar & Jesorka, Aldo (2015). Lipid Self-Spreading on Solid Substrates. IntechOpen. ISBN 978-953-51-2216-6. 36 p.
• Gözen, Irep (2013). The self–spreading double bilayer/Advances in Lipid Membrane Nanotechnology. Chalmers University of Technology. ISBN 978-91-7385-813-7. 110 p.

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• Thiyagaraja, Maivizhi & Gözen, Irep (2023). Thermoresponsive artificial cells. Biophysical Journal. ISSN 0006-3495. 122(3). doi: 10.1016/j.bpj.2022.11.1555.
• Balaban, Theodor Silviu; Gözen, Irep & Jesorka, Aldo (2016). The Nobel Prize for Chemistry 2016 highlights tiny synthetic machines. NBS-nytt. ISSN 0801-3535. p. 11–11.
• Schanke, Ingrid Jin & Gözen, Irep (2021). Transport within protocell-nanotube networks and its implications for communication of primitive cells. Universitetet i Oslo. Show summary

  Protocells are self-organized primitive compartments which are thought to be precursors to first living cells at the origin of life. Several different types of model protocell structures have been proposed; among them, the most recently identified protocell nanotube networks (PNNs). A PNN consists of tens of surface adhered lipid compartments physically connected via lipid nanotubes. In this work, it is shown for the first time using differential interference contrast (DIC) microscopy that the PNNs can successfully form by using lipid compositions that are completely label-free. It has also been shown via confocal microscopy that the lipid nanotubes allow the transport of fluorescently labeled water soluble compounds, RNA and DNA among the compartments in the network. By employing fluorescence recovery after photobleaching (FRAP) technique, the diffusive redistribution of encapsulated fluorophores among compartments that are not in direct contact with each other, has been depicted. Results show that the transportation of the compounds in PNNs is possible due to the interconnecting nanotubes, which is proposed as a plausible chemical communication and replication mechanism between the protocells on the early Earth.

• Köksal, Elif; Gözen, Irep; Lund, Reidar & Carlson, Andreas (2021). Surface-assisted formation and development of model protocells. Universitetet i Oslo, Det matematisk-naturvitenskapelige fakultet. ISSN 1501-7710. 2021(2401). Show summary

  By enabling formation and development of primitive cells, also known as protocells, solid surfaces might have played an important role in the emergence of life on Earth. Current theories on how life originated from primitive cells are based on the initial step, where a spherical capsule made from fat molecules formed spontaneously in water. This model explains the formation well, but not the following steps, such as growth, replication and division. In this PhD thesis, the possible involvement of solid surfaces in the formation and development of primitive cells has been investigated. Solid surfaces existed in the form of rocks and minerals on the early Earth. By bringing together three simple ingredients: solid surfaces, fat molecules and water, and observing their interactions by means of microscopy, a series of unique transformations has been identified. Examples of such behavior are the formation of protocell networks connected with lipid nanotubes, formation of subcompartments similar to organelles in modern cells, protocell fusion and colony formation. Nano-engineered artificial surfaces, as well as natural surfaces including glasses, minerals, rocks, and a meteorite fragment from Mars originating from the earliest crust of the planet were used in the studies.

• Spustova, Karolina; Gözen, Irep; Lund, Reidar & Carlson, Andreas (2021). Membranous Protocell Superstructures. Universitetet i Oslo, Det matematisk-naturvitenskapelige fakultet. ISSN 1501-7710. 2021(2467). Show summary

  How primitive cells emerged and developed on the early Earth remain an unsolved question. In contrast to the hypotheses that the first living cells evolved from a single ancestor, their development as colonies would provide numerous advantages, such as mechanical stability, enhancement of chemical reactions, and sharing of prebiotic molecules. In this PhD thesis, the formation and development of colony-like primitive cell structures and populations was investigated using three simple ingredients: water, fat molecules and solid surfaces. Transformation of fat agglomerates into large populations of structured primitive cells that are able to communicate and migrate, was observed under a light microscope. The role of solid surfaces which were abundant on the early Earth in form of rocks and minerals, was shown to be fundamental for these processes. These findings make us rethink the current theories on how life originated and show that solid surfaces might have played an important role in the emergence of primitive cells on the early Earth.

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