By the end of the course, students will be able to: Quantitative reasoning at the cellular scale ● Apply statistical mechanics and physical reasoning to estimate forces, timescales, and material parameters of cellular components. ● Derive and use key mechanical models — persistence length, flexural rigidity, Helfrich bending energy, entropic elasticity — to describe the behaviour of biopolymers and membranes. Mechanics of cellular components ● Describe the structure and mechanical function of cytoskeletal filaments, polymer networks, molecular motors, and lipid membranes, and explain how their properties arise from physical principles. ● Analyse how mechanical and thermodynamic constraints shape the design of biological machinery. Mechanobiology and cellular response ● Explain how cells sense and transmit mechanical forces, and connect these mechanisms to key mechanotransduction pathways and biological outcomes. Critical reading and independent thinking ● Interpret quantitative results from primary literature and experimental methods used to probe mechanical properties at the cell and tissue scale. ● Develop analytical problem-solving skills through structured exercises and case-based examples.

Students are expected to have a solid background in classical mechanics, thermodynamics, and introductory biology and chemistry. Familiarity with statistical mechanics and basic physical reasoning at the molecular scale is helpful. The lecturer will address specific background gaps through dedicated lectures.

Physical foundations ● Length, time, and force scales across biological systems ● Thermal energy (kT) and thermal fluctuations: why mechanics at the nanoscale defies macroscopic intuition ● Brownian motion, entropic forces, and the role of noise in biological design ● Dimensional analysis and order-of-magnitude estimation Mechanics of cellular components ● Biopolymers: persistence length, flexural rigidity, worm-like chain model, and entropic elasticity ● Cytoskeletal filaments: actin, microtubules, and intermediate filaments — mechanical properties, polymerisation dynamics, and structural roles ● Polymer network mechanics: elasticity of crosslinked and entangled gels ● Molecular motors: mechanochemical cycle, force-velocity relationships, and energetics of myosin, kinesin, and dynein ● Lipid membranes: bending rigidity, Helfrich energy, thermal shape fluctuations, and membrane tension ● The nucleus: chromatin mechanics and nuclear stiffness Mechanobiology: how cells sense and respond to force ● Extracellular matrix and cell-scale mechanics ● Mechanosensing: integrins, focal adhesions, and mechanosensitive ion channels ● Mechanotransduction: from force to biochemical signal ● Cell migration and locomotion: actin-driven motility and immune cell trafficking ● Neurobiomechanics: mechanics of neural tissue, axonal mechanics, and physical forces in neural function Applications and special topics Selected case studies and examples drawn from recent primary literature. Topics may include: tissue mechanics in health and disease (cancer, fibrosis, cardiovascular, neurological); emergent mechanical solutions in living organisms (insect biomechanics, spider silk, bioinspired materials). One dedicated topic explores the relationship between biological learning and artificial intelligence. Synaptic plasticity, immune memory, and swarm behaviour are learning implemented in physical, mechanical systems. Comparing these with their AI counterparts — gradient descent, classification algorithms, swarm optimisation — reveals what is gained and what is lost in the abstraction: understanding the mechanics of how living systems learn illuminates both biology and the principles behind its computational imitations.

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The course combines theoretical lectures with guided problem-solving sessions and case-based seminars to foster both conceptual understanding and practical reasoning skills. ● Lectures introduce and discuss the fundamental principles of cellular and tissue biomechanics, with frequent interdisciplinary references to physics, biology, chemistry, and engineering, and examples drawn from primary scientific literature. ● Classroom exercises involve analytical problem solving and order-of-magnitude estimation, applying theoretical models to real biological systems with guidance from the instructor. ● Case-based seminars explore specific biological or clinical phenomena through the lens of biomechanical reasoning — including analysis of landmark papers and discussion of open questions — to stimulate independent scientific thinking.

The teacher will provide all the course material (slides and lecture notes). Suggested textbooks: ● Boal, D.H., 2012. Mechanics of the Cell. 2nd ed. Cambridge University Press. ● Jacobs, C.R., Huang, H., Kwon, R.Y., 2012. Introduction to Cell Mechanics and Mechanobiology. Garland Science. ● Vogel, S., 2013. Comparative Biomechanics: Life's Physical World. 2nd ed. Princeton University Press. ● Phillips, R., Kondev, J., Theriot, J., Orme, N., 2012. Physical Biology of the Cell. 2nd ed. Garland Science. ● Howard, J., 2001. Mechanics of Motor Proteins and the Cytoskeleton. Sinauer Associates. ● Nelson, P., 2008. Biological Physics: Energy, Information, Life. W.H. Freeman. ● Mofrad, M.R.K., Kamm, R.D. (eds.), 2006. Cytoskeletal Mechanics: Models and Measurements in Cell Mechanics. Cambridge University Press.

Lecture slides; Lecture notes; Exercise with solutions ;

Exam: Written test;

The course is assessed through a written examination. The exam includes both theoretical questions and applied numerical problems. Questions are weighted based on complexity: 1 point for theoretical questions, and 2 to 3 points for applied problems. The maximum raw score is 34; a score above 31 corresponds to the top grade (30 cum laude).

In addition to the message sent by the online system, students with disabilities or Specific Learning Disorders (SLD) are invited to directly inform the professor in charge of the course about the special arrangements for the exam that have been agreed with the Special Needs Unit. The professor has to be informed at least one week before the beginning of the examination session in order to provide students with the most suitable arrangements for each specific type of exam.