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标题：Cell wall composition determines handedness reversal in helicoidal cellulose architectures of Pollia condensata fruits
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作者：Yin Chang ; Rox Middleton ; Yu Ogawa 等
期刊名称：Proceedings of the National Academy of Sciences
印刷版ISSN：0027-8424
电子版ISSN：1091-6490
出版年度：2021
卷号：118
期号：51
DOI：10.1073/pnas.2111723118
语种：English
出版社：The National Academy of Sciences of the United States of America
摘要：Significance Helicoidal architectures are widespread in nature; several species adopt this structure to produce brilliant colorations. Such chiral architectures are usually left-handed in plants, with the only exception found in the cell walls of epicarp cells of Pollia condensata, where both handednesses are observed. Here, we aim to understand the origin of handednesses by analyzing optical and mechanical responses of single cells. Surprisingly, we discover that left-handed and right-handed cells show different distributions of spectra and elasticity. We verified by using finite element analysis simulation that the elasticity of helicoids is sensitive to the ratio of cellulose/cell wall matrix. Our findings reveal that cell wall composition affects the helicoidal architectures, suggesting that chemical composition plays a role in morphogenesis of the chirality reversal. Chiral asymmetry is important in a wide variety of disciplines and occurs across length scales. While several natural chiral biomolecules exist only with single handedness, they can produce complex hierarchical structures with opposite chiralities. Understanding how the handedness is transferred from molecular to the macroscopic scales is far from trivial. An intriguing example is the transfer of the handedness of helicoidal organizations of cellulose microfibrils in plant cell walls. These cellulose helicoids produce structural colors if their dimension is comparable to the wavelength of visible light. All previously reported examples of a helicoidal structure in plants are left-handed except, remarkably, in the Pollia condensata fruit; both left- and right-handed helicoidal cell walls are found in neighboring cells of the same tissue. By simultaneously studying optical and mechanical responses of cells with different handednesses, we propose that the chirality of helicoids results from differences in cell wall composition. In detail, here we showed statistical substantiation of three different observations: 1) light reflected from right-handed cells is red shifted compared to light reflected from left-handed cells, 2) right-handed cells occur more rarely than left-handed ones, and 3) right-handed cells are located mainly in regions corresponding to interlocular divisions. Finally, 4) right-handed cells have an average lower elastic modulus compared to left-handed cells of the same color. Our findings, combined with mechanical simulation, suggest that the different chiralities of helicoids in the cell wall may result from different chemical composition, which strengthens previous hypotheses that hemicellulose might mediate the rotations of cellulose microfibrils.
关键词：helicoidal cell wall; chirality; structural colors; mechanical properties of plant cell wall; cellulose–hemicellulose interaction