Intracellular Mass Density Increase Is Accompanying but Not Sufficient for Stiffening and Growth Arrest of Yeast Cells

Many organisms, including yeast cells, bacteria, nematodes, and tardigrades, endure harsh environmental conditions, such as nutrient scarcity, or lack of water and energy for a remarkably long time. The rescue programs that these organisms launch upon encountering these adverse conditions include reprogramming their metabolism in order to enter a quiescent or dormant state in a controlled fashion. Reprogramming coincides with changes in the macromolecular architecture and changes in the physical and mechanical properties of the cells. However, the cellular mechanisms underlying the physical-mechanical changes remain enigmatic. Here, we induce metabolic arrest of yeast cells by lowering their intracellular pH. We then determine the differences in the intracellular mass density and stiffness of active and metabolically arrested cells using optical diffraction tomography (ODT) and atomic force microscopy (AFM). We show that an increased intracellular mass density is associated with an increase in stiffness when the growth of yeast is arrested. However, increasing the intracellular mass density alone is not sufficient for maintenance of the growth-arrested state in yeast cells. Our data suggest that the cytoplasm of metabolically arrested yeast displays characteristics of a solid. Our findings constitute a bridge between the mechanical behavior of the cytoplasm and the physical and chemical mechanisms of metabolically arrested cells with the ultimate aim of understanding dormant organisms

Piezoelectric Response of Polycrystalline Silicon-Doped Hafnium Oxide Thin Films Determined by Rapid Temperature Cycles

The in‐plane piezoelectric response of 20 nm thick Si‐doped HfO2 is examined by exploiting thermal expansion of the substrate upon rapid temperature cycling. The sample is heated locally by a deposited metal film, and the subsequently registered pyroelectric current is found to be frequency dependent in the observed range of 5 Hz to 35 kHz. While the intrinsic response remains constant, the secondary contribution can be switched off in the high‐frequency limit due to substrate clamping. As this secondary response is generated by thermal expansion and the piezoelectric effect, this allows for extraction of the corresponding in‐plane response. By comparing pyroelectric measurements in low‐ and high‐frequency limits, a piezoelectric coefficient d 31 of −11.5 pm V −1 is obtained, which is more than five times larger than that of AlN. The magnitude of piezoelectric response increases upon electric field cycling, which is associated with a transition from antiferroelectric‐like behavior to a purely ferroelectric polarization hysteresis. The hafnium oxide material system is proposed as a promising candidate for future CMOS compatible piezoelectric micro‐ and nano‐electromechanical systems (MEMS and NEMS)