Arman Toofani, Sepehr H. Eraghi, Ali Basti, Hamed Rajabi
Presenting a novel framework for sustainable and regenerative design and development is a fundamental future need. Here we argue that a new framework, referred to as complexity biomechanics, which can be used for holistic analysis and understanding of natural mechanical systems, is key to fulfilling this need. We also present a roadmap for the design and development of intelligent and complex engineering materials, mechanisms, structures, systems, and processes capable of automatic adaptation and self-organization in response to ever-changing environments. We apply complexity biomechanics to elucidate how the different structural components of a complex biological system as dragonfly wings, from ultrastructure of the cuticle, the constituting bio-composite material of the wing, to higher structural levels, collaboratively contribute to the functionality of the entire wing system. This framework not only proposes a paradigm shift in understanding and drawing inspiration from natural systems but also holds potential applications in various domains, including materials science and engineering, biomechanics, biomimetics, bionics, and engineering biology.
{"title":"Complexity biomechanics: a case study of dragonfly wing design from constituting composite material to higher structural levels","authors":"Arman Toofani, Sepehr H. Eraghi, Ali Basti, Hamed Rajabi","doi":"10.1098/rsfs.2023.0060","DOIUrl":"https://doi.org/10.1098/rsfs.2023.0060","url":null,"abstract":"<p>Presenting a novel framework for sustainable and regenerative design and development is a fundamental future need. Here we argue that a new framework, referred to as complexity biomechanics, which can be used for holistic analysis and understanding of natural mechanical systems, is key to fulfilling this need. We also present a roadmap for the design and development of intelligent and complex engineering materials, mechanisms, structures, systems, and processes capable of automatic adaptation and self-organization in response to ever-changing environments. We apply complexity biomechanics to elucidate how the different structural components of a complex biological system as dragonfly wings, from ultrastructure of the cuticle, the constituting bio-composite material of the wing, to higher structural levels, collaboratively contribute to the functionality of the entire wing system. This framework not only proposes a paradigm shift in understanding and drawing inspiration from natural systems but also holds potential applications in various domains, including materials science and engineering, biomechanics, biomimetics, bionics, and engineering biology.</p>","PeriodicalId":13795,"journal":{"name":"Interface Focus","volume":"89 1","pages":""},"PeriodicalIF":4.4,"publicationDate":"2024-04-12","publicationTypes":"Journal Article","fieldsOfStudy":null,"isOpenAccess":false,"openAccessPdf":"","citationCount":null,"resultStr":null,"platform":"Semanticscholar","paperid":"140575651","PeriodicalName":null,"FirstCategoryId":null,"ListUrlMain":null,"RegionNum":3,"RegionCategory":"生物学","ArticlePicture":[],"TitleCN":null,"AbstractTextCN":null,"PMCID":"","EPubDate":null,"PubModel":null,"JCR":null,"JCRName":null,"Score":null,"Total":0}
Elaine M. Bast, Natalie T. Marshall, Kendall O. Myers, Lucas W. Marsh, Martin Walschburger Hurtado, Peter A. Van Zandt, Matthew S. Lehnert
Insects have evolved unique structures that host a diversity of material and mechanical properties, and the mouthparts (proboscis) of butterflies and moths (Lepidoptera) are no exception. Here, we examined proboscis morphology and material properties from several previously unstudied moth lineages to determine if they relate to flower visiting and non-flower visiting feeding habits. Scanning electron microscopy and three-dimensional imaging were used to study proboscis morphology and assess surface roughness patterns on the galeal surface, respectively. Confocal laser scanning microscopy was used to study patterns of cuticular autofluorescence, which was quantified with colour analysis software. We found that moth proboscises display similar autofluorescent signals and morphological patterns in relation to feeding habits to those previously described for flower and non-flower visiting butterflies. The distal region of proboscises of non-flower visitors is brush-like for augmented capillarity and exhibited blue autofluorescence, indicating the possible presence of resilin and increased flexibility. Flower visitors have smoother proboscises and show red autofluorescence, an indicator of high sclerotization, which is adaptive for floral tube entry. We propose the lepidopteran proboscis as a model structure for understanding how insects have evolved a suite of morphological and material adaptations to overcome the challenges of acquiring fluids from diverse sources.
{"title":"Diverse material properties and morphology of moth proboscises relates to the feeding habits of some macromoth and other lepidopteran lineages","authors":"Elaine M. Bast, Natalie T. Marshall, Kendall O. Myers, Lucas W. Marsh, Martin Walschburger Hurtado, Peter A. Van Zandt, Matthew S. Lehnert","doi":"10.1098/rsfs.2023.0051","DOIUrl":"https://doi.org/10.1098/rsfs.2023.0051","url":null,"abstract":"<p>Insects have evolved unique structures that host a diversity of material and mechanical properties, and the mouthparts (proboscis) of butterflies and moths (Lepidoptera) are no exception. Here, we examined proboscis morphology and material properties from several previously unstudied moth lineages to determine if they relate to flower visiting and non-flower visiting feeding habits. Scanning electron microscopy and three-dimensional imaging were used to study proboscis morphology and assess surface roughness patterns on the galeal surface, respectively. Confocal laser scanning microscopy was used to study patterns of cuticular autofluorescence, which was quantified with colour analysis software. We found that moth proboscises display similar autofluorescent signals and morphological patterns in relation to feeding habits to those previously described for flower and non-flower visiting butterflies. The distal region of proboscises of non-flower visitors is brush-like for augmented capillarity and exhibited blue autofluorescence, indicating the possible presence of resilin and increased flexibility. Flower visitors have smoother proboscises and show red autofluorescence, an indicator of high sclerotization, which is adaptive for floral tube entry. We propose the lepidopteran proboscis as a model structure for understanding how insects have evolved a suite of morphological and material adaptations to overcome the challenges of acquiring fluids from diverse sources.</p>","PeriodicalId":13795,"journal":{"name":"Interface Focus","volume":"20 1","pages":""},"PeriodicalIF":4.4,"publicationDate":"2024-04-12","publicationTypes":"Journal Article","fieldsOfStudy":null,"isOpenAccess":false,"openAccessPdf":"","citationCount":null,"resultStr":null,"platform":"Semanticscholar","paperid":"140575657","PeriodicalName":null,"FirstCategoryId":null,"ListUrlMain":null,"RegionNum":3,"RegionCategory":"生物学","ArticlePicture":[],"TitleCN":null,"AbstractTextCN":null,"PMCID":"","EPubDate":null,"PubModel":null,"JCR":null,"JCRName":null,"Score":null,"Total":0}
Svenja Hackethal, Ellen Schulz-Kornas, Stanislav N. Gorb, Wencke Krings
Radular teeth have to cope with wear, when interacting with ingesta. In some molluscan taxa, wear-coping mechanisms, related to the incorporation of high contents of iron or silica, have been previously determined. For most species, particularly for those which possess radulae without such incorporations, wear-coping mechanisms are understudied. In the present study, we documented and characterized the wear on radular teeth in the model species Loligo vulgaris (Cephalopoda). By applying a range of methods, the elementary composition and mechanical properties of the teeth were described, to gain insight into mechanisms for coping with abrasion. It was found that the tooth regions that are prone to wear are harder and stiffer. Additionally, the surfaces interacting with the ingesta possessed a thin coating with high contents of silicon, probably reducing abrasion. The here presented data may serve as an example of systematic study of radular wear, in order to understand the relationship between the structure of radular teeth and their properties.
{"title":"Wear patterns of radular teeth in Loligo vulgaris (Cephalopoda; Mollusca) are related to their structure and mechanical properties","authors":"Svenja Hackethal, Ellen Schulz-Kornas, Stanislav N. Gorb, Wencke Krings","doi":"10.1098/rsfs.2023.0082","DOIUrl":"https://doi.org/10.1098/rsfs.2023.0082","url":null,"abstract":"<p>Radular teeth have to cope with wear, when interacting with ingesta. In some molluscan taxa, wear-coping mechanisms, related to the incorporation of high contents of iron or silica, have been previously determined. For most species, particularly for those which possess radulae without such incorporations, wear-coping mechanisms are understudied. In the present study, we documented and characterized the wear on radular teeth in the model species <i>Loligo vulgaris</i> (Cephalopoda). By applying a range of methods, the elementary composition and mechanical properties of the teeth were described, to gain insight into mechanisms for coping with abrasion. It was found that the tooth regions that are prone to wear are harder and stiffer. Additionally, the surfaces interacting with the ingesta possessed a thin coating with high contents of silicon, probably reducing abrasion. The here presented data may serve as an example of systematic study of radular wear, in order to understand the relationship between the structure of radular teeth and their properties.</p>","PeriodicalId":13795,"journal":{"name":"Interface Focus","volume":"15 1","pages":""},"PeriodicalIF":4.4,"publicationDate":"2024-04-12","publicationTypes":"Journal Article","fieldsOfStudy":null,"isOpenAccess":false,"openAccessPdf":"","citationCount":null,"resultStr":null,"platform":"Semanticscholar","paperid":"140589862","PeriodicalName":null,"FirstCategoryId":null,"ListUrlMain":null,"RegionNum":3,"RegionCategory":"生物学","ArticlePicture":[],"TitleCN":null,"AbstractTextCN":null,"PMCID":"","EPubDate":null,"PubModel":null,"JCR":null,"JCRName":null,"Score":null,"Total":0}
Valentin Birkenfeld, Stanislav N. Gorb, Wencke Krings
Leafcutter ant colonies are divided into castes with the individuals performing different tasks, based mostly on size. With the mandibles, the small minims care for the brood or the fungus, whereas the larger minors and mediae cut and transport plant material, with the ant size positively related to the material size. The mechanical properties and composition of the mandible cuticle have been previously tested in the soldiers as the largest caste, revealing that the cutting edges contained high contents of the cross-linking transition metal zinc (Zn). With regard to the smaller castes, no data are present. To study how the mandible size and function relates to its mechanical properties, we here tested the mandibles of minims, minors and mediae by nanoindentation. We found that the hardness (H) and Young's modulus (E) values increased with increasing ant size and that the mandible cutting edges in each caste have the highest H- and E-values. To gain insight into the origins of these properties, we characterized the elemental composition by energy-dispersive X-ray analysis, revealing that minors and mediae possessed higher content of Zn in the cutting edges in contrast to the minims containing significantly less Zn. This shows, that Zn content relates to higher mechanical property values. Additionally, it shows that all of these parameters can differ within a single species.
切叶蚁群分为不同的种群,个体主要根据大小执行不同的任务。体型较小的小蚁用下颚照顾蚁巢或真菌,而体型较大的小蚁和中蚁则切割和运输植物材料,蚂蚁的体型与材料的大小呈正相关。以前曾对作为最大种姓的兵蚁的下颚角质层的机械性能和成分进行过测试,结果表明切割边缘含有较多的交联过渡金属锌(Zn)。至于较小的种姓,目前还没有相关数据。为了研究下颌骨的大小和功能与其机械性能之间的关系,我们在此通过纳米压痕法测试了微小种、微小种和中等种的下颌骨。我们发现,硬度(H)和杨氏模量(E)值随着蚂蚁体型的增大而增大,而且每个种姓的下颚切削刃都具有最高的 H 值和 E 值。为了深入了解这些特性的来源,我们通过能量色散 X 射线分析来确定元素组成的特征,结果表明,未成年蚂蚁和中型蚂蚁的切削刃中锌含量较高,而未成年蚂蚁的锌含量明显较低。这表明,锌含量与较高的机械性能值有关。此外,它还表明所有这些参数在同一品种中也会有所不同。
{"title":"Mandible elemental composition and mechanical properties from distinct castes of the leafcutter ant Atta laevigata (Attini; Formicidae)","authors":"Valentin Birkenfeld, Stanislav N. Gorb, Wencke Krings","doi":"10.1098/rsfs.2023.0048","DOIUrl":"https://doi.org/10.1098/rsfs.2023.0048","url":null,"abstract":"<p>Leafcutter ant colonies are divided into castes with the individuals performing different tasks, based mostly on size. With the mandibles, the small minims care for the brood or the fungus, whereas the larger minors and mediae cut and transport plant material, with the ant size positively related to the material size. The mechanical properties and composition of the mandible cuticle have been previously tested in the soldiers as the largest caste, revealing that the cutting edges contained high contents of the cross-linking transition metal zinc (Zn). With regard to the smaller castes, no data are present. To study how the mandible size and function relates to its mechanical properties, we here tested the mandibles of minims, minors and mediae by nanoindentation. We found that the hardness (H) and Young's modulus (E) values increased with increasing ant size and that the mandible cutting edges in each caste have the highest H- and E-values. To gain insight into the origins of these properties, we characterized the elemental composition by energy-dispersive X-ray analysis, revealing that minors and mediae possessed higher content of Zn in the cutting edges in contrast to the minims containing significantly less Zn. This shows, that Zn content relates to higher mechanical property values. Additionally, it shows that all of these parameters can differ within a single species.</p>","PeriodicalId":13795,"journal":{"name":"Interface Focus","volume":"30 1","pages":""},"PeriodicalIF":4.4,"publicationDate":"2024-04-12","publicationTypes":"Journal Article","fieldsOfStudy":null,"isOpenAccess":false,"openAccessPdf":"","citationCount":null,"resultStr":null,"platform":"Semanticscholar","paperid":"140575487","PeriodicalName":null,"FirstCategoryId":null,"ListUrlMain":null,"RegionNum":3,"RegionCategory":"生物学","ArticlePicture":[],"TitleCN":null,"AbstractTextCN":null,"PMCID":"","EPubDate":null,"PubModel":null,"JCR":null,"JCRName":null,"Score":null,"Total":0}
Anika Preuss, Esther Appel, Stanislav N. Gorb, Sebastian Büsse
The arthropod cuticle offers strength, protection, and lightweight. Due to its limit in expandability, arthropods have to moult periodically to grow. While moulting is beneficial in terms of parasite or toxin control, growth and adaptation to environmental conditions, it costs energy and leaves the soft animal's body vulnerable to injuries and desiccation directly after ecdysis. To investigate the temporal change in sclerotization and pigmentation during and after ecdysis, we combined macrophotography, confocal laser scanning microscopy, scanning electron microscopy and histological sectioning. We analysed the tarsal and mandibular cuticle of the blue emperor dragonfly to compare the progress of tanning for structures that are functionally involved during emergence (tarsus/tarsal claws) with structures whose functionality is required much later (mandibles). Our results show that: (i) the tanning of the tarsal and mandibular cuticle increases during emergence; (ii) the tarsal cuticle tans faster than the mandibular cuticle; (iii) the mandibles tan faster on the aboral than on the oral side; and (iv) both the exo- and the endocuticle are tanned. The change in the cuticle composition of the tarsal and mandibular cuticle reflects the demand for higher mechanical stability of these body parts when holding on to the substrate during emergence and during first walking or hunting attempts.
{"title":"Tanning of the tarsal and mandibular cuticle in adult Anax imperator (Insecta: Odonata) during the emergence sequence","authors":"Anika Preuss, Esther Appel, Stanislav N. Gorb, Sebastian Büsse","doi":"10.1098/rsfs.2023.0076","DOIUrl":"https://doi.org/10.1098/rsfs.2023.0076","url":null,"abstract":"<p>The arthropod cuticle offers strength, protection, and lightweight. Due to its limit in expandability, arthropods have to moult periodically to grow. While moulting is beneficial in terms of parasite or toxin control, growth and adaptation to environmental conditions, it costs energy and leaves the soft animal's body vulnerable to injuries and desiccation directly after ecdysis. To investigate the temporal change in sclerotization and pigmentation during and after ecdysis, we combined macrophotography, confocal laser scanning microscopy, scanning electron microscopy and histological sectioning. We analysed the tarsal and mandibular cuticle of the blue emperor dragonfly to compare the progress of tanning for structures that are functionally involved during emergence (tarsus/tarsal claws) with structures whose functionality is required much later (mandibles). Our results show that: (i) the tanning of the tarsal and mandibular cuticle increases during emergence; (ii) the tarsal cuticle tans faster than the mandibular cuticle; (iii) the mandibles tan faster on the aboral than on the oral side; and (iv) both the exo- and the endocuticle are tanned. The change in the cuticle composition of the tarsal and mandibular cuticle reflects the demand for higher mechanical stability of these body parts when holding on to the substrate during emergence and during first walking or hunting attempts.</p>","PeriodicalId":13795,"journal":{"name":"Interface Focus","volume":"52 1","pages":""},"PeriodicalIF":4.4,"publicationDate":"2024-04-12","publicationTypes":"Journal Article","fieldsOfStudy":null,"isOpenAccess":false,"openAccessPdf":"","citationCount":null,"resultStr":null,"platform":"Semanticscholar","paperid":"140589863","PeriodicalName":null,"FirstCategoryId":null,"ListUrlMain":null,"RegionNum":3,"RegionCategory":"生物学","ArticlePicture":[],"TitleCN":null,"AbstractTextCN":null,"PMCID":"","EPubDate":null,"PubModel":null,"JCR":null,"JCRName":null,"Score":null,"Total":0}
The mandibles of the desert locust Schistocerca gregaria (Forsskål, 1775) are digger-shovel-shaped mouthparts that are part of the locust's exoskeleton formed by the insect cuticle. The cuticle is a polymer–fibre composite, which supports, encases and protects the entire body. Mandibles experience heavy loading and wear due to direct contact with hard and abrasive food, just like teeth, their mineralized analogues in vertebrates. With dual-energy X-ray tomography, we image well-defined regions of zinc (Zn)-enriched cuticle at the mandible cutting edges and quantify the Zn concentrations in these regions. Zn is known to increase stiffness, hardness and wear resistance of the otherwise purely polymeric insect cuticle. In S. gregaria, the position of the Zn-enriched cutting-edge regions relative to one another suggests that the mandibles form a scissor-like cutting tool, which sharpens itself as the mouthparts shear past one another during feeding. Comparing the architecture of these purely polymeric mandibles with the mineralized incisors of rodents, we find fundamental design differences in cutting-tool structure and performance. Locusts' scissors and rodents’ carving knives perform different functions, because they act on food that differs significantly in properties and shape: softer, sheet-like material in the case of locusts and harder bulk material in the case of rodents.
沙漠蝗虫 Schistocerca gregaria(Forsskål,1775 年)的下颚是挖掘铲形口器,是由昆虫角质层形成的蝗虫外骨骼的一部分。角质层是一种聚合物-纤维复合材料,支撑、包裹并保护整个身体。下颚由于直接接触坚硬和磨蚀性的食物而承受着巨大的负荷和磨损,就像脊椎动物的矿化类似物--牙齿一样。通过双能 X 射线断层扫描,我们对下颌骨切削边缘富锌(Zn)角质层的明确区域进行了成像,并对这些区域的锌浓度进行了量化。众所周知,锌能增加昆虫角质层的硬度、刚度和耐磨性。在 S. gregaria 中,富含锌的切割边缘区域的相对位置表明,下颚形成了一个类似剪刀的切割工具,在进食过程中,当口器相互剪切时,该工具会变得锋利。将这些纯聚合物下颚的结构与啮齿动物的矿化门齿进行比较,我们发现两者在切割工具的结构和性能方面存在根本性的设计差异。蝗虫的剪刀和啮齿动物的刻刀发挥着不同的功能,因为它们作用的食物在性质和形状上有很大的不同:蝗虫的食物是较软的片状材料,而啮齿动物的食物则是较硬的块状材料。
{"title":"Desert locusts (Schistocerca gregaria) feed with self-sharpening, scissor-like mandibles","authors":"Ulrike G. K. Wegst, Peter Cloetens, Oliver Betz","doi":"10.1098/rsfs.2023.0069","DOIUrl":"https://doi.org/10.1098/rsfs.2023.0069","url":null,"abstract":"<p>The mandibles of the desert locust <i>Schistocerca gregaria</i> (Forsskål, 1775) are digger-shovel-shaped mouthparts that are part of the locust's exoskeleton formed by the insect cuticle. The cuticle is a polymer–fibre composite, which supports, encases and protects the entire body. Mandibles experience heavy loading and wear due to direct contact with hard and abrasive food, just like teeth, their mineralized analogues in vertebrates. With dual-energy X-ray tomography, we image well-defined regions of zinc (Zn)-enriched cuticle at the mandible cutting edges and quantify the Zn concentrations in these regions. Zn is known to increase stiffness, hardness and wear resistance of the otherwise purely polymeric insect cuticle. In <i>S. gregaria,</i> the position of the Zn-enriched cutting-edge regions relative to one another suggests that the mandibles form a scissor-like cutting tool, which sharpens itself as the mouthparts shear past one another during feeding. Comparing the architecture of these purely polymeric mandibles with the mineralized incisors of rodents, we find fundamental design differences in cutting-tool structure and performance. Locusts' scissors and rodents’ carving knives perform different functions, because they act on food that differs significantly in properties and shape: softer, sheet-like material in the case of locusts and harder bulk material in the case of rodents.</p>","PeriodicalId":13795,"journal":{"name":"Interface Focus","volume":"72 1","pages":""},"PeriodicalIF":4.4,"publicationDate":"2024-04-12","publicationTypes":"Journal Article","fieldsOfStudy":null,"isOpenAccess":false,"openAccessPdf":"","citationCount":null,"resultStr":null,"platform":"Semanticscholar","paperid":"140602609","PeriodicalName":null,"FirstCategoryId":null,"ListUrlMain":null,"RegionNum":3,"RegionCategory":"生物学","ArticlePicture":[],"TitleCN":null,"AbstractTextCN":null,"PMCID":"","EPubDate":null,"PubModel":null,"JCR":null,"JCRName":null,"Score":null,"Total":0}
Locust Locusta migratoria exhibits remarkable aerial performances, relying predominantly on its hind wings that generate most of lift and thrust for flight. The mechanical properties of the cross-veins determine the deformation of the hind wing, which greatly affect the aerodynamic performance of flapping flight. However, whether the mechanical behaviours of the locust cross-veins change with loading rate is still unknown. In this study, cross-veins in four physiological regions (anterior–medial, anterior–lateral, posterior–medial and posterior–lateral) of the hind wing from adult locusts were investigated using uniaxial tensile test, stress relaxation test and fluorescence microscopy. It was found that the cross-veins were a type of viscoelastic material (including rate-independent elastic modulus and obvious stress relaxation). The cross-veins in the two anterior regions of the hind wing had significantly higher elastic moduli and higher ultimate tensile stress than those of its two posterior regions. This difference might be attributed to different resilin distribution patterns in the cross-veins. These findings furnish new insights into the mechanical characteristics of the locust cross-veins, which might deepen our understanding of the aerodynamic mechanisms of locust flapping flight.
{"title":"The mechanical properties of different cross-veins in the hind wing of locust Locusta migratoria under uniaxial tensile and stress relaxation tests","authors":"Yizun Zhou, Linxin Bai, Chao Wan","doi":"10.1098/rsfs.2023.0068","DOIUrl":"https://doi.org/10.1098/rsfs.2023.0068","url":null,"abstract":"<p>Locust <i>Locusta migratoria</i> exhibits remarkable aerial performances, relying predominantly on its hind wings that generate most of lift and thrust for flight. The mechanical properties of the cross-veins determine the deformation of the hind wing, which greatly affect the aerodynamic performance of flapping flight. However, whether the mechanical behaviours of the locust cross-veins change with loading rate is still unknown. In this study, cross-veins in four physiological regions (anterior–medial, anterior–lateral, posterior–medial and posterior–lateral) of the hind wing from adult locusts were investigated using uniaxial tensile test, stress relaxation test and fluorescence microscopy. It was found that the cross-veins were a type of viscoelastic material (including rate-independent elastic modulus and obvious stress relaxation). The cross-veins in the two anterior regions of the hind wing had significantly higher elastic moduli and higher ultimate tensile stress than those of its two posterior regions. This difference might be attributed to different resilin distribution patterns in the cross-veins. These findings furnish new insights into the mechanical characteristics of the locust cross-veins, which might deepen our understanding of the aerodynamic mechanisms of locust flapping flight.</p>","PeriodicalId":13795,"journal":{"name":"Interface Focus","volume":"70 1","pages":""},"PeriodicalIF":4.4,"publicationDate":"2024-04-12","publicationTypes":"Journal Article","fieldsOfStudy":null,"isOpenAccess":false,"openAccessPdf":"","citationCount":null,"resultStr":null,"platform":"Semanticscholar","paperid":"140575630","PeriodicalName":null,"FirstCategoryId":null,"ListUrlMain":null,"RegionNum":3,"RegionCategory":"生物学","ArticlePicture":[],"TitleCN":null,"AbstractTextCN":null,"PMCID":"","EPubDate":null,"PubModel":null,"JCR":null,"JCRName":null,"Score":null,"Total":0}
Daniela E. Winkler, Hitomi Seike, Shinji Nagata, Mugino O. Kubo
Animals have evolved diverse comminuting tools. While vertebrates possess mineralized teeth, insect mandibles often bear metal-inclusion-hardened serrated cusps. Microscopic dental enamel wear (microwear) is known to be caused by contact with ingesta. To test if insect mandible microwear is also diet-dependent, we kept newly moulted adult two-spotted crickets (Gryllus bimaculatus) for four weeks on alfalfa-based rodent pellets with and without added mineral abrasives (loess, quartz, volcanic ash). Six crickets per diet were examined after 1, 3, 7, 14, 21 and 28 days. All diets induced progressive mandible wear, affecting specific locations along the distal tooth cusps differently. The depth of furrows increased on most abrasive-containing diets until day 21, while wear mark complexity increased from day 1 to 3 and 14 to 21. After 28 days, these parameter values for large volcanic ash and large quartz diets significantly exceeded those for the control diet. These results are comparable to observations from guinea pig feeding experiments with the same diets. Cricket mandible wear was affected by all abrasives. Notably, large volcanic ash and large quartz induced the deepest, most complex lesions, akin to observations in guinea pigs. This suggests a universal wear process, supporting that microwear analyses are suitable for inferring invertebrate diets.
{"title":"Mandible microwear texture analysis of crickets raised on diets of different abrasiveness reveals universality of diet-induced wear","authors":"Daniela E. Winkler, Hitomi Seike, Shinji Nagata, Mugino O. Kubo","doi":"10.1098/rsfs.2023.0065","DOIUrl":"https://doi.org/10.1098/rsfs.2023.0065","url":null,"abstract":"<p>Animals have evolved diverse comminuting tools. While vertebrates possess mineralized teeth, insect mandibles often bear metal-inclusion-hardened serrated cusps. Microscopic dental enamel wear (microwear) is known to be caused by contact with ingesta. To test if insect mandible microwear is also diet-dependent, we kept newly moulted adult two-spotted crickets (<i>Gryllus bimaculatus</i>) for four weeks on alfalfa-based rodent pellets with and without added mineral abrasives (loess, quartz, volcanic ash). Six crickets per diet were examined after 1, 3, 7, 14, 21 and 28 days. All diets induced progressive mandible wear, affecting specific locations along the distal tooth cusps differently. The depth of furrows increased on most abrasive-containing diets until day 21, while wear mark complexity increased from day 1 to 3 and 14 to 21. After 28 days, these parameter values for large volcanic ash and large quartz diets significantly exceeded those for the control diet. These results are comparable to observations from guinea pig feeding experiments with the same diets. Cricket mandible wear was affected by all abrasives. Notably, large volcanic ash and large quartz induced the deepest, most complex lesions, akin to observations in guinea pigs. This suggests a universal wear process, supporting that microwear analyses are suitable for inferring invertebrate diets.</p>","PeriodicalId":13795,"journal":{"name":"Interface Focus","volume":"20 1","pages":""},"PeriodicalIF":4.4,"publicationDate":"2024-04-12","publicationTypes":"Journal Article","fieldsOfStudy":null,"isOpenAccess":false,"openAccessPdf":"","citationCount":null,"resultStr":null,"platform":"Semanticscholar","paperid":"140575631","PeriodicalName":null,"FirstCategoryId":null,"ListUrlMain":null,"RegionNum":3,"RegionCategory":"生物学","ArticlePicture":[],"TitleCN":null,"AbstractTextCN":null,"PMCID":"","EPubDate":null,"PubModel":null,"JCR":null,"JCRName":null,"Score":null,"Total":0}
Cristian L. Klunk, Michael Heethoff, Jörg U. Hammel, Stanislav N. Gorb, Wencke Krings
Mandible morphology has an essential role in biting performance, but the mandible cuticle can have regional differences in its mechanical properties. The effects of such a heterogeneous distribution of cuticle material properties in the mandible responses to biting loading are still poorly explored in chewing insects. Here, we tested the mechanical properties of mandibles of the ant species Formica cunicularia by nanoindentation and investigated the effects of the cuticular variation in Young's modulus (E) under bite loading with finite-element analysis (FEA). The masticatory margin of the mandible, which interacts with the food, was the hardest and stiffest region. To unravel the origins of the mechanical property gradients, we characterized the elemental composition by energy-dispersive X-ray spectroscopy. The masticatory margin possessed high proportions of Cu and Zn. When incorporated into the FEA, variation in E effectively changed mandible stress patterns, leading to a relatively higher concentration of stresses in the stiffer mandibular regions and leaving the softer mandible blade with relatively lower stress. Our results demonstrated the relevance of cuticle E heterogeneity in mandibles under bite loading, suggesting that the accumulation of transition metals such as Cu and Zn has a relevant correlation with the mechanical characteristics in F. cunicularia mandibles.
下颚形态对咬合性能起着至关重要的作用,但下颚角质层的机械性能可能存在区域差异。在咀嚼昆虫中,这种角质层材料特性的异质性分布对下颚咬合负荷反应的影响还没有得到充分的探讨。在此,我们通过纳米压痕法测试了蚂蚁物种Formica cunicularia下颚的机械性能,并利用有限元分析(FEA)研究了咬合加载下杨氏模量(E)的角质层变化的影响。下颌骨的咀嚼边缘与食物相互作用,是最硬和最僵硬的区域。为了揭示机械性能梯度的起源,我们通过能量色散 X 射线光谱分析确定了元素组成。咀嚼边缘含有较高比例的铜和锌。在纳入有限元分析后,E 的变化有效地改变了下颌骨的应力模式,导致较硬的下颌骨区域应力相对较高,而较软的下颌骨叶片应力相对较低。我们的研究结果表明了咬合负荷下下颌骨角质层 E 异质性的相关性,表明铜和锌等过渡金属的积累与 F. cunicularia 下颌骨的机械特性具有相关性。
{"title":"Mechanical and elemental characterization of ant mandibles: consequences for bite mechanics","authors":"Cristian L. Klunk, Michael Heethoff, Jörg U. Hammel, Stanislav N. Gorb, Wencke Krings","doi":"10.1098/rsfs.2023.0056","DOIUrl":"https://doi.org/10.1098/rsfs.2023.0056","url":null,"abstract":"<p>Mandible morphology has an essential role in biting performance, but the mandible cuticle can have regional differences in its mechanical properties. The effects of such a heterogeneous distribution of cuticle material properties in the mandible responses to biting loading are still poorly explored in chewing insects. Here, we tested the mechanical properties of mandibles of the ant species <i>Formica cunicularia</i> by nanoindentation and investigated the effects of the cuticular variation in Young's modulus (E) under bite loading with finite-element analysis (FEA). The masticatory margin of the mandible, which interacts with the food, was the hardest and stiffest region. To unravel the origins of the mechanical property gradients, we characterized the elemental composition by energy-dispersive X-ray spectroscopy. The masticatory margin possessed high proportions of Cu and Zn. When incorporated into the FEA, variation in E effectively changed mandible stress patterns, leading to a relatively higher concentration of stresses in the stiffer mandibular regions and leaving the softer mandible blade with relatively lower stress. Our results demonstrated the relevance of cuticle E heterogeneity in mandibles under bite loading, suggesting that the accumulation of transition metals such as Cu and Zn has a relevant correlation with the mechanical characteristics in <i>F</i>. <i>cunicularia</i> mandibles.</p>","PeriodicalId":13795,"journal":{"name":"Interface Focus","volume":"41 1","pages":""},"PeriodicalIF":4.4,"publicationDate":"2024-04-12","publicationTypes":"Journal Article","fieldsOfStudy":null,"isOpenAccess":false,"openAccessPdf":"","citationCount":null,"resultStr":null,"platform":"Semanticscholar","paperid":"140575632","PeriodicalName":null,"FirstCategoryId":null,"ListUrlMain":null,"RegionNum":3,"RegionCategory":"生物学","ArticlePicture":[],"TitleCN":null,"AbstractTextCN":null,"PMCID":"","EPubDate":null,"PubModel":null,"JCR":null,"JCRName":null,"Score":null,"Total":0}
The crustacean cuticle is a biological composite material consisting of chitin–protein fibres in a mineralized matrix. Recent research has revealed a surprising range of fibre architectures and mineral compositions of crustacean skeletal structures adapted to various mechanical demands. It is becoming increasingly clear that the organic fibres in the cuticle may be organized in patterns differing from the standard twisted plywood model. Observed fibre architectures in protruding skeletal structures include longitudinal and circular parallel fibre arrays. Skeletal minerals often include calcium phosphates in addition to calcium carbonates. Furthermore, skeletal properties are affected by protein cross-linking, which replaces mineralization as a stiffening mechanism in some structures. Several common structural motifs, such as the stiffening of the outer skeletal layers, the incorporation of non-mineralized cuticle in exposed structures, and interchanging layers of parallel fibres and the twisted plywood structure, can be identified in skeletal elements with similar functions. These evolutionary solutions have the potential for biomimetic applications, particularly as manufacturing technologies advance. To make use of this potential, we need to understand the processes behind the formation of the crustacean exoskeleton and determine which features are truly adaptive and worth mimicking.
{"title":"Structural diversity of crustacean exoskeletons and its implications for biomimetics","authors":"Miloš Vittori","doi":"10.1098/rsfs.2023.0075","DOIUrl":"https://doi.org/10.1098/rsfs.2023.0075","url":null,"abstract":"<p>The crustacean cuticle is a biological composite material consisting of chitin–protein fibres in a mineralized matrix. Recent research has revealed a surprising range of fibre architectures and mineral compositions of crustacean skeletal structures adapted to various mechanical demands. It is becoming increasingly clear that the organic fibres in the cuticle may be organized in patterns differing from the standard twisted plywood model. Observed fibre architectures in protruding skeletal structures include longitudinal and circular parallel fibre arrays. Skeletal minerals often include calcium phosphates in addition to calcium carbonates. Furthermore, skeletal properties are affected by protein cross-linking, which replaces mineralization as a stiffening mechanism in some structures. Several common structural motifs, such as the stiffening of the outer skeletal layers, the incorporation of non-mineralized cuticle in exposed structures, and interchanging layers of parallel fibres and the twisted plywood structure, can be identified in skeletal elements with similar functions. These evolutionary solutions have the potential for biomimetic applications, particularly as manufacturing technologies advance. To make use of this potential, we need to understand the processes behind the formation of the crustacean exoskeleton and determine which features are truly adaptive and worth mimicking.</p>","PeriodicalId":13795,"journal":{"name":"Interface Focus","volume":"24 1","pages":""},"PeriodicalIF":4.4,"publicationDate":"2024-04-12","publicationTypes":"Journal Article","fieldsOfStudy":null,"isOpenAccess":false,"openAccessPdf":"","citationCount":null,"resultStr":null,"platform":"Semanticscholar","paperid":"140590003","PeriodicalName":null,"FirstCategoryId":null,"ListUrlMain":null,"RegionNum":3,"RegionCategory":"生物学","ArticlePicture":[],"TitleCN":null,"AbstractTextCN":null,"PMCID":"","EPubDate":null,"PubModel":null,"JCR":null,"JCRName":null,"Score":null,"Total":0}