title: Mechanical characterization of outer epidermal middle lamella of onion under tensile loading.
abstract: •
  
  
  
  The cells in plant tissue are joined together by a distinct layer called the middle lamella (ML). Understanding the mechanical properties of the ML is crucial in studying how tissue-level mechanical properties emerge from the subcellular-level mechanical properties. However, the nanoscale size of the ML presents formidable challenges to its characterization as a separate layer. Consequently, the mechanical properties of the ML under tensile loading are as yet unknown.•
  
  
  
  Here, we characterize the ML from a subcellular sample excised from two adjacent cells and composed of two wall fragments and a single line of ML in between. Two techniques, cryotome sectioning and milling with a focused ion beam, were used to prepare ML samples, and tensile experiments were performed using microelectromechanical system (MEMS) tensile testing devices.•
  
  
  
  Our test results showed that even at a subcellular scale, the ML appears to be stronger than the wall fragments. There was also evidence that the ML attached at the corner of cells more strongly than at the rest of the contact area. The contribution of the additional ML contact area was estimated to be 40.6 MPa. Wall fragment samples containing an ML layer were also significantly stronger (p < 0.05) than the wall fragments without an ML layer.•
  
  
  
  The tensile properties of the ML might not have a major impact on the tissue-scale mechanical properties. This conclusion calls for further study of the ML, including characterization under shear loading conditions and elucidation of the contributions of other extracellular parameters, such as cell size and shape, to the overall tissue-level mechanical response.
authors.affiliation: Department of Agricultural and Biological Engineering, 229 Agricultural Engineering Building, Pennsylvania State University, University Park, Pennsylvania 16802 USA.
authors.forename: M Shafayet
authors.initials: MS
authors.lastname: Zamil
authors.affiliation: Department of Agricultural and Biological Engineering, 229 Agricultural Engineering Building, Pennsylvania State University, University Park, Pennsylvania 16802 USA.
authors.forename: Hojae
authors.initials: H
authors.lastname: Yi
authors.affiliation: Department of Agricultural and Biological Engineering, 229 Agricultural Engineering Building, Pennsylvania State University, University Park, Pennsylvania 16802 USA.
authors.forename: Virendra M
authors.initials: VM
authors.lastname: Puri
mesh_terms: D001696:Biomechanical Phenomena; D055617:Micro-Electrical-Mechanical Systems; D019697:Onions / Q000502:physiology*; D019441:Plant Epidermis / Q000502:physiology*; D013314:Stress, Mechanical; D013718:Tensile Strength
keywords: Allium cepa; Amaryllidaceae; MEMS; mechanical properties; middle lamella; onion; plant cell wall; tensile test