Avoidance of Catastrophic Structural Failure as an Evolutionary Constraint: Biomechanics of the Acorn Weevil Rostrum
The acorn weevil (Curculio Linnaeus, 1758) rostrum (snout) exhibits remarkable flexibility and toughness derived from the microarchitecture of its exoskeleton. Modifications to the composite profile of the rostral cuticle that simultaneously enhance the flexibility and toughness of the distal portio...
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Published in: | Advanced materials (Weinheim) Vol. 31; no. 41; pp. e1903526 - n/a |
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Abstract | The acorn weevil (Curculio Linnaeus, 1758) rostrum (snout) exhibits remarkable flexibility and toughness derived from the microarchitecture of its exoskeleton. Modifications to the composite profile of the rostral cuticle that simultaneously enhance the flexibility and toughness of the distal portion of the snout are characterized. Using classical laminate plate theory, the effect of these modifications on the elastic behavior of the exoskeleton is estimated. It is shown that the tensile behavior of the rostrum across six Curculio species with high morphological variation correlates with changes in the relative layer thicknesses and orientation angles of layers in the exoskeleton. Accordingly, increased endocuticle thickness is strongly correlated with increased tensile strength. Rostrum stiffness is shown to be inversely correlated with work of fracture; thus allowing a highly curved rostrum to completely straighten without structural damage. Finally, exocuticle rich invaginations of the occipital sutures are identified both as a likely site of crack initiation in tensile failure and as a source of morphological constraint on the evolution of the rostrum in Curculio weevils. It is concluded that avoidance of catastrophic structural failure, as initiated in these sutures under tension, is the driving selective pressure in the evolution of the female Curculio rostrum.
The effect of heterogeneous laminate microstructure on cuticle strength in the acorn weevil rostrum is investigated. The results indicate that increased endocuticle thickness and advantageous fiber orientation near the rostral apex allows the snout to be repeatedly subjected to extreme bending without harm. These modifications mitigate the risk of catastrophic structural failure through a tradeoff between stiffness and toughness of the laminate. |
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AbstractList | The acorn weevil (Curculio Linnaeus, 1758) rostrum (snout) exhibits remarkable flexibility and toughness derived from the microarchitecture of its exoskeleton. Modifications to the composite profile of the rostral cuticle that simultaneously enhance the flexibility and toughness of the distal portion of the snout are characterized. Using classical laminate plate theory, the effect of these modifications on the elastic behavior of the exoskeleton is estimated. It is shown that the tensile behavior of the rostrum across six Curculio species with high morphological variation correlates with changes in the relative layer thicknesses and orientation angles of layers in the exoskeleton. Accordingly, increased endocuticle thickness is strongly correlated with increased tensile strength. Rostrum stiffness is shown to be inversely correlated with work of fracture; thus allowing a highly curved rostrum to completely straighten without structural damage. Finally, exocuticle rich invaginations of the occipital sutures are identified both as a likely site of crack initiation in tensile failure and as a source of morphological constraint on the evolution of the rostrum in Curculio weevils. It is concluded that avoidance of catastrophic structural failure, as initiated in these sutures under tension, is the driving selective pressure in the evolution of the female Curculio rostrum. The acorn weevil (Curculio Linnaeus, 1758) rostrum (snout) exhibits remarkable flexibility and toughness derived from the microarchitecture of its exoskeleton. Modifications to the composite profile of the rostral cuticle that simultaneously enhance the flexibility and toughness of the distal portion of the snout are characterized. Using classical laminate plate theory, the effect of these modifications on the elastic behavior of the exoskeleton is estimated. It is shown that the tensile behavior of the rostrum across six Curculio species with high morphological variation correlates with changes in the relative layer thicknesses and orientation angles of layers in the exoskeleton. Accordingly, increased endocuticle thickness is strongly correlated with increased tensile strength. Rostrum stiffness is shown to be inversely correlated with work of fracture; thus allowing a highly curved rostrum to completely straighten without structural damage. Finally, exocuticle rich invaginations of the occipital sutures are identified both as a likely site of crack initiation in tensile failure and as a source of morphological constraint on the evolution of the rostrum in Curculio weevils. It is concluded that avoidance of catastrophic structural failure, as initiated in these sutures under tension, is the driving selective pressure in the evolution of the female Curculio rostrum. The effect of heterogeneous laminate microstructure on cuticle strength in the acorn weevil rostrum is investigated. The results indicate that increased endocuticle thickness and advantageous fiber orientation near the rostral apex allows the snout to be repeatedly subjected to extreme bending without harm. These modifications mitigate the risk of catastrophic structural failure through a tradeoff between stiffness and toughness of the laminate. The acorn weevil ( Curculio Linnaeus, 1758) rostrum (snout) exhibits remarkable flexibility and toughness derived from the microarchitecture of its exoskeleton. Modifications to the composite profile of the rostral cuticle that simultaneously enhance the flexibility and toughness of the distal portion of the snout are characterized. Using classical laminate plate theory, the effect of these modifications on the elastic behavior of the exoskeleton is estimated. It is shown that the tensile behavior of the rostrum across six Curculio species with high morphological variation correlates with changes in the relative layer thicknesses and orientation angles of layers in the exoskeleton. Accordingly, increased endocuticle thickness is strongly correlated with increased tensile strength. Rostrum stiffness is shown to be inversely correlated with work of fracture; thus allowing a highly curved rostrum to completely straighten without structural damage. Finally, exocuticle rich invaginations of the occipital sutures are identified both as a likely site of crack initiation in tensile failure and as a source of morphological constraint on the evolution of the rostrum in Curculio weevils. It is concluded that avoidance of catastrophic structural failure, as initiated in these sutures under tension, is the driving selective pressure in the evolution of the female Curculio rostrum. |
Author | Jansen, Michael A. Franz, Nico M. Chawla, Nikhilesh Williams, Jason |
Author_xml | – sequence: 1 givenname: Michael A. orcidid: 0000-0001-6289-6541 surname: Jansen fullname: Jansen, Michael A. email: majanse1@asu.edu organization: Arizona State University – sequence: 2 givenname: Jason surname: Williams fullname: Williams, Jason organization: Energy, and Transport Arizona State University – sequence: 3 givenname: Nikhilesh surname: Chawla fullname: Chawla, Nikhilesh organization: Energy, and Transport Arizona State University – sequence: 4 givenname: Nico M. surname: Franz fullname: Franz, Nico M. organization: Arizona State University |
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Cites_doi | 10.1016/j.actbio.2011.04.004 10.1126/sciadv.aao5469 10.1080/19768354.2017.1330764 10.1007/BF00561216 10.1242/jcs.21.1.73 10.1073/pnas.1222787110 10.1073/pnas.1409585111 10.1098/rspa.2017.0607 10.1201/b12409 10.1038/nmat4309 10.1098/rstb.1942.0003 10.1016/j.matchar.2016.05.022 10.1007/978-1-349-16673-2 10.1111/jbi.12733 10.1557/jmr.2009.0409 10.1111/j.0014-3820.2004.tb00483.x 10.1016/j.jmbbm.2010.09.015 10.1016/j.asd.2004.05.006 10.1086/498277 10.4028/www.scientific.net/AMR.581-582.1133 10.1242/jeb.083824 10.5109/2677 10.1086/284325 10.1016/j.asd.2015.10.002 10.1016/S0040-8166(72)80042-9 10.1016/j.ympev.2004.02.007 10.1016/0022-2836(80)90156-4 10.1242/jeb.068221 10.1007/s00442-011-2036-7 10.1098/rspb.2014.0696 10.1016/j.jsb.2016.05.007 10.1111/j.2041-210X.2010.00044.x 10.1002/adma.200902019 10.1016/0020-7322(92)90004-7 |
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Keywords | tensile testing fractography acorn weevils exoskeletons |
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Snippet | The acorn weevil (Curculio Linnaeus, 1758) rostrum (snout) exhibits remarkable flexibility and toughness derived from the microarchitecture of its exoskeleton.... The acorn weevil ( Curculio Linnaeus, 1758) rostrum (snout) exhibits remarkable flexibility and toughness derived from the microarchitecture of its... |
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SubjectTerms | acorn weevils Animals Avoidance Biological Evolution Biomechanical Phenomena Biomechanics Catastrophic events Computer architecture Correlation Crack initiation Elastic Modulus Elasticity Evolution Exoskeletons Female Flexibility fractography Fracture toughness Laminates Materials science Mechanical Phenomena Morphology Plate theory Stiffness Structural damage Structural failure Tensile Strength tensile testing Thickness Weevils - anatomy & histology Weevils - physiology |
Title | Avoidance of Catastrophic Structural Failure as an Evolutionary Constraint: Biomechanics of the Acorn Weevil Rostrum |
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