Jon Barnes - Academia.edu (original) (raw)
Papers by Jon Barnes
Journal of Experimental Biology, 2003
SUMMARYFiddler crabs Uca rapax are central-place foragers, making feeding excursions of up to sev... more SUMMARYFiddler crabs Uca rapax are central-place foragers, making feeding excursions of up to several meters from their burrows. This study investigates the sources of directional and distance information used by these crabs when returning to their burrows. We tested the spatial frame of reference(egocentric or exocentric), and the source of spatial information (idiothetic or allothetic) used during homing. We also tested which components of their locomotion they integrated (only voluntary, or voluntary plus reflexive).Fiddler crabs in their natural mudflat habitat were passively rotated during normal foraging behavior using experimenter-controlled disks, before they returned home. Crabs resisted passive rotations on the disk by counter-rotating when the disk turned, which was a compensatory response to unintended movement. Crabs were usually situated eccentrically on the disk,and therefore were also subjected to a translation when the disk rotated. No crab actively compensated for ...
Journal of Experimental Biology, 2008
SUMMARY Knowledge of both surface structure and physical properties such as stiffness and elastic... more SUMMARY Knowledge of both surface structure and physical properties such as stiffness and elasticity are essential to understanding any adhesive system. In this study of an adhesion surface in the tree frog, Litoria caerulea White, a variety of techniques including atomic force microscopy were used to investigate the microstructure and properties of an epithelium that adheres through wet adhesion. Litoria toe pads consist of a hexagonal array of flat-topped epithelial cells, separated by mucus-filled channels. Under an atomic force microscope, this `flat' surface is highly structured at the nanoscale, consisting of a tightly packed array of columnar nanopillars (described as hemidesmosomes by previous authors), 326±84 nm in diameter, each of which possesses a central dimple 8±4 nm in depth. In fixed tissue (transmission electron microscopy), the nanopillars are approximately as tall as they are broad. At the gross anatomical level, larger toe pads may be subdivided into medial a...
Journal of Morphology, 2013
Cryo-scanning electron microscopy (cryo-SEM) and atomic force microscopy (AFM) offer new avenues ... more Cryo-scanning electron microscopy (cryo-SEM) and atomic force microscopy (AFM) offer new avenues for the study of the morphology of tree frog adhesive toe pads. Using these techniques, we compare toe pad microstructure in two distantly related species of tree frog, Litoria caerulea, White (Hylidae) and Rhacophorus prominanus, Smith (Rhacophoridae), in which the toe pads are considered to be convergent. AFM demonstrates the extraordinary similarity of both surface microstructures (largely hexagonal epithelial cells surrounded by deep channels) and nanostructures (an array of nanopillars, ca. 350 nm in diameter, all with a small dimple at the apex). The cryo-SEM studies examined the distribution of the fibrillar cytoskeleton within the different layers of the stratified toe pad epithelium, demonstrating that the cytoskeletal elements (keratin tonofilaments) that lie at an angle to the surface are relatively poorly developed in L. caerulea, clearly so in comparison to R. prominanus. Cryo-SEM also enabled the visualization of the fluid layer that is critical to a toe pad's adhesive function. This was achieved by examination of the frozen fluid residues left behind after removal of a toe within the cryo-SEM's experimental chamber. Such 'toeprints' demonstrated the presence of a wedge of fluid surrounding each toe pad, as well as fluid filling the channels that surround each epithelial cell. Cryo-SEM was used to examine epithelial cell shape. In a sample of 582 cells, 59.5% were hexagonal, the remainder being mainly pentagonal (23.1%) or heptagonal (16.1%). The distribution of differently-shaped cells was not random, but was not associated with either pad curvature or the distribution of mucous pores that provide fluid for the frogs' wet adhesion mechanism. Our main finding, the great similarity of toe pad structure in these two species, has important implications for biomimetics, for such convergent evolution suggests a good starting point for attempts to develop adhesives that will function in wet conditions.
Journal of Comparative Physiology A, 2006
This allometric study of adhesion in 15 Trinidadian tree frog species investigates how relationsh... more This allometric study of adhesion in 15 Trinidadian tree frog species investigates how relationships between length, area and mass limit the ability of adult frog species of different sizes to adhere to inclined and overhanging surfaces. Our experiments show that hylid frogs possess an area-based wet adhesive system in which larger species are lighter than expected from isometry and adhere better than expected from their toe pad area. However, in spite of these adaptations, larger species adhere less well than smaller species. In addition to these adhesive forces, tree frogs also generate significant shear forces that scale with mass, suggesting that they are frictional forces. Toe pads detach by peeling and frogs have g Viscosity h F Angle of fall h S Angle of slip h 1 and h 2 Contact angles between fluid and adjoining surfaces l Coefficient of friction
Journal of Experimental Biology, 2003
SUMMARY Fiddler crabs Uca rapax are central-place foragers, making feeding excursions of up to 2 ... more SUMMARY Fiddler crabs Uca rapax are central-place foragers, making feeding excursions of up to 2 m from their burrows. We describe the natural feeding excursions of path-integrating fiddler crabs and analyze their paths for signs of significant systematic or random navigation errors. No signs of any systematic errors are evident. Random errors are small, probably due to a combination of the short length and low sinuosity of the foraging paths, as well as the fiddler crabs' unique method of locomotion that allows them to remain oriented to their burrows throughout the foraging path and to minimize large body turns. We further examined the extent to which their body orientation during foraging (transverse body axis pointing more or less towards home) accurately represented their stored home vector. By examining sequences of fast escape, we have shown that crabs can correct for deviations of their transverse body axis from home during their escape path. Thus their stored home vecto...
Journal of Experimental Biology, 2011
SUMMARYThis paper describes a search for neurones sensitive to optic flow in the visual system of... more SUMMARYThis paper describes a search for neurones sensitive to optic flow in the visual system of the shore crab Carcinus maenas using a procedure developed from that of Krapp and Hengstenberg. This involved determining local motion sensitivity and its directional selectivity at many points within the neurone's receptive field and plotting the results on a map. Our results showed that local preferred directions of motion are independent of velocity, stimulus shape and type of motion (circular or linear). Global response maps thus clearly represent real properties of the neurones' receptive fields. Using this method, we have discovered two families of interneurones sensitive to translational optic flow. The first family has its terminal arborisations in the lobula of the optic lobe, the second family in the medulla. The response maps of the lobula neurones (which appear to be monostratified lobular giant neurones) show a clear focus of expansion centred on or just above the h...
Journal of Comparative Physiology A, 2011
Previous work using an atomic force microscope in nanoindenter mode indicated that the outer, 10-... more Previous work using an atomic force microscope in nanoindenter mode indicated that the outer, 10-to 15-lm thick, keratinised layer of tree frog toe pads has a modulus of elasticity equivalent to silicone rubber (5-15 MPa) (Scholz et al. 2009), but gave no information on the physical properties of deeper structures. In this study, micro-indentation is used to measure the stiffness of whole toe pads of the tree frog, Litoria caerulea. We show here that tree frog toe pads are amongst the softest of biological structures (effective elastic modulus 4-25 kPa), and that they exhibit a gradient of stiffness, being stiffest on the outside. This stiffness gradient results from the presence of a dense network of capillaries lying beneath the pad epidermis, which probably has a shock absorbing function. Additionally, we compare the physical properties (elastic modulus, work of adhesion, pull-off force) of the toe pads of immature and adult frogs. Keywords Tree frog Á Adhesion Á Micro-indentation Á Effective elastic modulus Á Litoria caerulea Pablo Perez Goodwyn, our friend and collaborator, tragically died in a swimming accident during the summer of 2009. This paper is dedicated to his memory.
Journal of Experimental Biology
1. When a crayfish walks forward, its legs are not loaded equally. The third pair is the one exer... more 1. When a crayfish walks forward, its legs are not loaded equally. The third pair is the one exerting the largest vertically directed force, and the fourth legs produce the major part of the propulsive force. 2. Chronic recordings from a CSD2 nerve were made in crayfish walking on a treadmill. The activity of CSD2 rises during the stance phase. The activity peak at the beginning of the power stroke is especially pronounced in leg 4, which was studied in detail. The average spike frequency of CSD2 recordings during the first part of the power stroke is positively correlated with the initial increase in force produced by the leg. This suggests that CSD2 responds especially well to rapid load changes. 3. When a wire was fixed around a fourth leg such that it depressed the patch of compliant cuticle associated with CSD2, the activity of the sense organ was elevated throughout the entire step cycle, and the time between leg 4 touching the ground and leg 3 lifting off was more variable th...
Comparative animal mucomics: Inspiration for functional materials from ubiquitous and understudied biopolymers
ACS Biomaterials Science & Engineering
The functions of secreted animal mucuses are remarkably diverse and include lubricants, wet adhes... more The functions of secreted animal mucuses are remarkably diverse and include lubricants, wet adhesives, protective barriers, and mineralizing agents. Although present in all animals, many open questions related to the hierarchical architectures, material properties, and genetics of mucus remain. Here, we summarize what is known about secreted mucus structure, describe the work of research groups throughout the world who are investigating various animal mucuses, and relate how these studies are revealing new mucus properties and the relationships between mucus hierarchical structure and hydrogel function. Finally, we call for a more systematic approach to studying animal mucuses so that data sets can be compared, omics-style, to address unanswered questions in the emerging field of mucomics. One major result that we anticipate from these efforts is design rules for creating new materials that are inspired by the structures and functions of animal mucuses.
A force-measuring and behaviour-recording system consisting of 24 individual 3D force plates for the study of single limb forces in climbing animals on a quasi-cylindrical tower
Bioinspiration & Biomimetics
This study describes the design of a new force measuring array with a quasi-cylindrical surface f... more This study describes the design of a new force measuring array with a quasi-cylindrical surface for measuring the 3D ground reaction forces of animals climbing on a surface with high curvature. This force-measuring array was assembled from 24 individual 3D force sensors, each with a resolution at the millinewton (mN) level, which were installed from top to bottom in four columns and six rows, with sensors in neighbouring columns staggered in height. Three cameras were used to simultaneously record the climbing behaviours of animals (in these experiments tree frogs) on the cylinder-like force measuring array. We were thus able to simultaneously record the ground reaction forces of each of the four limbs of tree frogs (here six individuals of the Chinese gliding or flying frog, Rhacophorus dennysi, with forelimb spans in the range 163-201 mm) climbing or descending both smooth and rough surfaces on a quasi-cylindrical structure with an overall diameter of 79 mm. We describe the design and calibration of the individual force sensors, their installation and arrangement on the quasi-cylindrical climbing tower, the recording of ground reaction forces and climbing behaviour, data transformations necessitated by the angular relationship of neighbouring sensors, and data processing using MATLAB scripts. Additionally, we present preliminary data on the use of a clamping grip by climbing frogs and the existence of small pull-off forces that aid toe-pad detachment at the end of each locomotor stance phase.
The Journal of Experimental Biology
The adhesive mechanisms of climbing animals have become an important research topic because of th... more The adhesive mechanisms of climbing animals have become an important research topic because of their biomimetic implications. We examined the climbing abilities of hylid tree frogs on vertical cylinders of differing diameter and surface roughness to investigate the relative roles of adduction forces (gripping) and adhesion. Tree frogs adhere using their toe pads and subarticular tubercles, the adhesive joint being fluid-filled. Our hypothesis was that, on an effectively flat surface (adduction forces on the largest 120 mm diameter cylinder were insufficient to allow climbing), adhesion would effectively be the only means by which tree frogs could climb, but on the two smaller diameter cylinders (44 mm and 13 mm), frogs could additionally utilise adduction forces by gripping the cylinder either with their limbs outstretched or by grasping around the cylinder with their digits, respectively. The frogs’ performance would also depend on whether the surfaces were smooth (easy to adhere t...
Beilstein journal of nanotechnology, 2016
Tree frogs need to adhere to surfaces of various roughnesses in their natural habitats; these inc... more Tree frogs need to adhere to surfaces of various roughnesses in their natural habitats; these include bark, leaves and rocks. Rough surfaces can alter the effectiveness of their toe pads, due to factors such as a change of real contact area and abrasion of the pad epithelium. Here, we tested the effect of surface roughness on the attachment abilities of the tree frog Litoria caerulea. This was done by testing shear and adhesive forces on artificial surfaces with controlled roughness, both on single toe pads and whole animal scales. It was shown that frogs can stick 2-3 times better on small scale roughnesses (3-6 µm asperities), producing higher adhesive and frictional forces, but relatively poorly on the larger scale roughnesses tested (58.5-562.5 µm asperities). Our experiments suggested that, on such surfaces, the pads secrete insufficient fluid to fill the space under the pad, leaving air pockets that would significantly reduce the Laplace pressure component of capillarity. Ther...
Visual Systems: Neural Mechanisms and Visual Behavior
Crustacean Experimental Systems in Neurobiology, 2002
Page 221. Visual Systems: Neural Mechanisms and Visual Behavior Raymon M. Glantz'and W. Jon.... more Page 221. Visual Systems: Neural Mechanisms and Visual Behavior Raymon M. Glantz'and W. Jon. P. Barnes 2 'Department of Biochemistry and Cell Biology, Rice University, Houston, Texas, USA 2Division of Environmental ...
Adhesion in Wet Environments: Frogs
Encyclopedia of Nanotechnology, 2012
SUMMARY 1. When a crayfish walks forward, its legs are not loaded equally. The third pair is the ... more SUMMARY 1. When a crayfish walks forward, its legs are not loaded equally. The third pair is the one exerting the largest vertically directed force, and the fourth legs produce the major part of the propulsive force. 2. Chronic recordings from a CSD2 nerve were made in crayfish walking on a treadmill. The activity of CSD2 rises during the stance phase. The activity peak at the beginning of the power stroke is especially pronounced in leg 4, which was studied in detail. The average spike frequency of CSD2 recordings during the first part of the power stroke is positively correlated with the initial increase in force produced by the leg. This suggests that CSD2 responds especially well to rapid load changes. 3. When a wire was fixed around a fourth leg such that it depressed the patch of compliant cuticle associated with CSD2, the activity of the sense organ was elevated throughout the entire step cycle, and the time between leg 4 touching the ground and leg 3 lifting off was more var...
Effects of light intensity and pattern contrast on the ability of the land crab, Cardisoma guanhumi, to separate optic flow-field components
Visual Neuroscience, 2004
Using a novel suite of computer-generated visual stimuli that mimicked components of optic flow, ... more Using a novel suite of computer-generated visual stimuli that mimicked components of optic flow, the visual responses of the tropical land crab, Cardisoma guanhumi, were investigated. We show that crabs are normally successful in distinguishing the rotational and translational components of the optic flow field, showing strong optokinetic responses to the former but not the latter. This ability was not dependant on the orientation of the crab, occurring both in “forwards-walking” and “sideways-walking” configurations. However, under conditions of low overall light intensity and/or low object/background contrast, the separation mechanism shows partial failure causing the crab to generate compensatory eye movements to translation, particularly in response to low-frequency (low-velocity) stimuli. Using this discovery, we then tested the ability of crabs to separate rotational and translational components in a combined rotation/translation flow field under different conditions. We demon...
Local mechanisms for the separation of optic flow-field components in the land crab, Cardisoma guanhumi: A role for motion parallax?
Visual Neuroscience, 2004
Although a number of global mechanisms have been proposed over the years that explain how crabs m... more Although a number of global mechanisms have been proposed over the years that explain how crabs might separate the rotational and translational components of their optic flow field, there has been no evidence to date that local mechanisms such as motion parallax are used in this separation. We describe here a study that takes advantage of a recently developed suite of computer-generated visual stimuli that creates a three-dimensional world surrounding the crab in which we can simulate translational and rotational optic flow. We show that, while motion parallax is not the only mechanism used in flow-field separation, it does play a role in the recognition of translational optic flow fields in that, under conditions of low overall light intensity and low contrast ratio when crabs find the distinction between rotation and translation harder, smaller eye movements occur in response to translation when motion parallax cues are present than when they are absent. Thus, motion parallax is o...
Journal of Experimental Biology, 2012
SUMMARY Tree frogs use adhesive toe pads for climbing on a variety of surfaces. They rely on wet ... more SUMMARY Tree frogs use adhesive toe pads for climbing on a variety of surfaces. They rely on wet adhesion, which is aided by the secretion of mucus. In nature, the pads will undoubtedly get contaminated regularly through usage, but appear to maintain their stickiness over time. Here, we show in two experiments that the toe pads of White's tree frogs (Litoria caerulea) quickly recover from contamination through a self-cleaning mechanism. We compared adhesive forces prior to and after contamination of (1) the whole animal on a rotatable platform and (2) individual toe pads in restrained frogs mimicking individual steps using a motorised stage. In both cases, the adhesive forces recovered after a few steps but this took significantly longer in single toe pad experiments from restrained frogs, showing that use of the pads increases recovery. We propose that both shear movements and a ‘flushing’ effect of the secreted mucus play an important role in shedding particles/contaminants.
Adhesion and allometry from metamorphosis to maturation in hylid tree frogs: a sticky problem
Journal of Zoology, 2006
Journal of Experimental Biology, 2003
SUMMARYFiddler crabs Uca rapax are central-place foragers, making feeding excursions of up to sev... more SUMMARYFiddler crabs Uca rapax are central-place foragers, making feeding excursions of up to several meters from their burrows. This study investigates the sources of directional and distance information used by these crabs when returning to their burrows. We tested the spatial frame of reference(egocentric or exocentric), and the source of spatial information (idiothetic or allothetic) used during homing. We also tested which components of their locomotion they integrated (only voluntary, or voluntary plus reflexive).Fiddler crabs in their natural mudflat habitat were passively rotated during normal foraging behavior using experimenter-controlled disks, before they returned home. Crabs resisted passive rotations on the disk by counter-rotating when the disk turned, which was a compensatory response to unintended movement. Crabs were usually situated eccentrically on the disk,and therefore were also subjected to a translation when the disk rotated. No crab actively compensated for ...
Journal of Experimental Biology, 2008
SUMMARY Knowledge of both surface structure and physical properties such as stiffness and elastic... more SUMMARY Knowledge of both surface structure and physical properties such as stiffness and elasticity are essential to understanding any adhesive system. In this study of an adhesion surface in the tree frog, Litoria caerulea White, a variety of techniques including atomic force microscopy were used to investigate the microstructure and properties of an epithelium that adheres through wet adhesion. Litoria toe pads consist of a hexagonal array of flat-topped epithelial cells, separated by mucus-filled channels. Under an atomic force microscope, this `flat' surface is highly structured at the nanoscale, consisting of a tightly packed array of columnar nanopillars (described as hemidesmosomes by previous authors), 326±84 nm in diameter, each of which possesses a central dimple 8±4 nm in depth. In fixed tissue (transmission electron microscopy), the nanopillars are approximately as tall as they are broad. At the gross anatomical level, larger toe pads may be subdivided into medial a...
Journal of Morphology, 2013
Cryo-scanning electron microscopy (cryo-SEM) and atomic force microscopy (AFM) offer new avenues ... more Cryo-scanning electron microscopy (cryo-SEM) and atomic force microscopy (AFM) offer new avenues for the study of the morphology of tree frog adhesive toe pads. Using these techniques, we compare toe pad microstructure in two distantly related species of tree frog, Litoria caerulea, White (Hylidae) and Rhacophorus prominanus, Smith (Rhacophoridae), in which the toe pads are considered to be convergent. AFM demonstrates the extraordinary similarity of both surface microstructures (largely hexagonal epithelial cells surrounded by deep channels) and nanostructures (an array of nanopillars, ca. 350 nm in diameter, all with a small dimple at the apex). The cryo-SEM studies examined the distribution of the fibrillar cytoskeleton within the different layers of the stratified toe pad epithelium, demonstrating that the cytoskeletal elements (keratin tonofilaments) that lie at an angle to the surface are relatively poorly developed in L. caerulea, clearly so in comparison to R. prominanus. Cryo-SEM also enabled the visualization of the fluid layer that is critical to a toe pad's adhesive function. This was achieved by examination of the frozen fluid residues left behind after removal of a toe within the cryo-SEM's experimental chamber. Such 'toeprints' demonstrated the presence of a wedge of fluid surrounding each toe pad, as well as fluid filling the channels that surround each epithelial cell. Cryo-SEM was used to examine epithelial cell shape. In a sample of 582 cells, 59.5% were hexagonal, the remainder being mainly pentagonal (23.1%) or heptagonal (16.1%). The distribution of differently-shaped cells was not random, but was not associated with either pad curvature or the distribution of mucous pores that provide fluid for the frogs' wet adhesion mechanism. Our main finding, the great similarity of toe pad structure in these two species, has important implications for biomimetics, for such convergent evolution suggests a good starting point for attempts to develop adhesives that will function in wet conditions.
Journal of Comparative Physiology A, 2006
This allometric study of adhesion in 15 Trinidadian tree frog species investigates how relationsh... more This allometric study of adhesion in 15 Trinidadian tree frog species investigates how relationships between length, area and mass limit the ability of adult frog species of different sizes to adhere to inclined and overhanging surfaces. Our experiments show that hylid frogs possess an area-based wet adhesive system in which larger species are lighter than expected from isometry and adhere better than expected from their toe pad area. However, in spite of these adaptations, larger species adhere less well than smaller species. In addition to these adhesive forces, tree frogs also generate significant shear forces that scale with mass, suggesting that they are frictional forces. Toe pads detach by peeling and frogs have g Viscosity h F Angle of fall h S Angle of slip h 1 and h 2 Contact angles between fluid and adjoining surfaces l Coefficient of friction
Journal of Experimental Biology, 2003
SUMMARY Fiddler crabs Uca rapax are central-place foragers, making feeding excursions of up to 2 ... more SUMMARY Fiddler crabs Uca rapax are central-place foragers, making feeding excursions of up to 2 m from their burrows. We describe the natural feeding excursions of path-integrating fiddler crabs and analyze their paths for signs of significant systematic or random navigation errors. No signs of any systematic errors are evident. Random errors are small, probably due to a combination of the short length and low sinuosity of the foraging paths, as well as the fiddler crabs' unique method of locomotion that allows them to remain oriented to their burrows throughout the foraging path and to minimize large body turns. We further examined the extent to which their body orientation during foraging (transverse body axis pointing more or less towards home) accurately represented their stored home vector. By examining sequences of fast escape, we have shown that crabs can correct for deviations of their transverse body axis from home during their escape path. Thus their stored home vecto...
Journal of Experimental Biology, 2011
SUMMARYThis paper describes a search for neurones sensitive to optic flow in the visual system of... more SUMMARYThis paper describes a search for neurones sensitive to optic flow in the visual system of the shore crab Carcinus maenas using a procedure developed from that of Krapp and Hengstenberg. This involved determining local motion sensitivity and its directional selectivity at many points within the neurone's receptive field and plotting the results on a map. Our results showed that local preferred directions of motion are independent of velocity, stimulus shape and type of motion (circular or linear). Global response maps thus clearly represent real properties of the neurones' receptive fields. Using this method, we have discovered two families of interneurones sensitive to translational optic flow. The first family has its terminal arborisations in the lobula of the optic lobe, the second family in the medulla. The response maps of the lobula neurones (which appear to be monostratified lobular giant neurones) show a clear focus of expansion centred on or just above the h...
Journal of Comparative Physiology A, 2011
Previous work using an atomic force microscope in nanoindenter mode indicated that the outer, 10-... more Previous work using an atomic force microscope in nanoindenter mode indicated that the outer, 10-to 15-lm thick, keratinised layer of tree frog toe pads has a modulus of elasticity equivalent to silicone rubber (5-15 MPa) (Scholz et al. 2009), but gave no information on the physical properties of deeper structures. In this study, micro-indentation is used to measure the stiffness of whole toe pads of the tree frog, Litoria caerulea. We show here that tree frog toe pads are amongst the softest of biological structures (effective elastic modulus 4-25 kPa), and that they exhibit a gradient of stiffness, being stiffest on the outside. This stiffness gradient results from the presence of a dense network of capillaries lying beneath the pad epidermis, which probably has a shock absorbing function. Additionally, we compare the physical properties (elastic modulus, work of adhesion, pull-off force) of the toe pads of immature and adult frogs. Keywords Tree frog Á Adhesion Á Micro-indentation Á Effective elastic modulus Á Litoria caerulea Pablo Perez Goodwyn, our friend and collaborator, tragically died in a swimming accident during the summer of 2009. This paper is dedicated to his memory.
Journal of Experimental Biology
1. When a crayfish walks forward, its legs are not loaded equally. The third pair is the one exer... more 1. When a crayfish walks forward, its legs are not loaded equally. The third pair is the one exerting the largest vertically directed force, and the fourth legs produce the major part of the propulsive force. 2. Chronic recordings from a CSD2 nerve were made in crayfish walking on a treadmill. The activity of CSD2 rises during the stance phase. The activity peak at the beginning of the power stroke is especially pronounced in leg 4, which was studied in detail. The average spike frequency of CSD2 recordings during the first part of the power stroke is positively correlated with the initial increase in force produced by the leg. This suggests that CSD2 responds especially well to rapid load changes. 3. When a wire was fixed around a fourth leg such that it depressed the patch of compliant cuticle associated with CSD2, the activity of the sense organ was elevated throughout the entire step cycle, and the time between leg 4 touching the ground and leg 3 lifting off was more variable th...
Comparative animal mucomics: Inspiration for functional materials from ubiquitous and understudied biopolymers
ACS Biomaterials Science & Engineering
The functions of secreted animal mucuses are remarkably diverse and include lubricants, wet adhes... more The functions of secreted animal mucuses are remarkably diverse and include lubricants, wet adhesives, protective barriers, and mineralizing agents. Although present in all animals, many open questions related to the hierarchical architectures, material properties, and genetics of mucus remain. Here, we summarize what is known about secreted mucus structure, describe the work of research groups throughout the world who are investigating various animal mucuses, and relate how these studies are revealing new mucus properties and the relationships between mucus hierarchical structure and hydrogel function. Finally, we call for a more systematic approach to studying animal mucuses so that data sets can be compared, omics-style, to address unanswered questions in the emerging field of mucomics. One major result that we anticipate from these efforts is design rules for creating new materials that are inspired by the structures and functions of animal mucuses.
A force-measuring and behaviour-recording system consisting of 24 individual 3D force plates for the study of single limb forces in climbing animals on a quasi-cylindrical tower
Bioinspiration & Biomimetics
This study describes the design of a new force measuring array with a quasi-cylindrical surface f... more This study describes the design of a new force measuring array with a quasi-cylindrical surface for measuring the 3D ground reaction forces of animals climbing on a surface with high curvature. This force-measuring array was assembled from 24 individual 3D force sensors, each with a resolution at the millinewton (mN) level, which were installed from top to bottom in four columns and six rows, with sensors in neighbouring columns staggered in height. Three cameras were used to simultaneously record the climbing behaviours of animals (in these experiments tree frogs) on the cylinder-like force measuring array. We were thus able to simultaneously record the ground reaction forces of each of the four limbs of tree frogs (here six individuals of the Chinese gliding or flying frog, Rhacophorus dennysi, with forelimb spans in the range 163-201 mm) climbing or descending both smooth and rough surfaces on a quasi-cylindrical structure with an overall diameter of 79 mm. We describe the design and calibration of the individual force sensors, their installation and arrangement on the quasi-cylindrical climbing tower, the recording of ground reaction forces and climbing behaviour, data transformations necessitated by the angular relationship of neighbouring sensors, and data processing using MATLAB scripts. Additionally, we present preliminary data on the use of a clamping grip by climbing frogs and the existence of small pull-off forces that aid toe-pad detachment at the end of each locomotor stance phase.
The Journal of Experimental Biology
The adhesive mechanisms of climbing animals have become an important research topic because of th... more The adhesive mechanisms of climbing animals have become an important research topic because of their biomimetic implications. We examined the climbing abilities of hylid tree frogs on vertical cylinders of differing diameter and surface roughness to investigate the relative roles of adduction forces (gripping) and adhesion. Tree frogs adhere using their toe pads and subarticular tubercles, the adhesive joint being fluid-filled. Our hypothesis was that, on an effectively flat surface (adduction forces on the largest 120 mm diameter cylinder were insufficient to allow climbing), adhesion would effectively be the only means by which tree frogs could climb, but on the two smaller diameter cylinders (44 mm and 13 mm), frogs could additionally utilise adduction forces by gripping the cylinder either with their limbs outstretched or by grasping around the cylinder with their digits, respectively. The frogs’ performance would also depend on whether the surfaces were smooth (easy to adhere t...
Beilstein journal of nanotechnology, 2016
Tree frogs need to adhere to surfaces of various roughnesses in their natural habitats; these inc... more Tree frogs need to adhere to surfaces of various roughnesses in their natural habitats; these include bark, leaves and rocks. Rough surfaces can alter the effectiveness of their toe pads, due to factors such as a change of real contact area and abrasion of the pad epithelium. Here, we tested the effect of surface roughness on the attachment abilities of the tree frog Litoria caerulea. This was done by testing shear and adhesive forces on artificial surfaces with controlled roughness, both on single toe pads and whole animal scales. It was shown that frogs can stick 2-3 times better on small scale roughnesses (3-6 µm asperities), producing higher adhesive and frictional forces, but relatively poorly on the larger scale roughnesses tested (58.5-562.5 µm asperities). Our experiments suggested that, on such surfaces, the pads secrete insufficient fluid to fill the space under the pad, leaving air pockets that would significantly reduce the Laplace pressure component of capillarity. Ther...
Visual Systems: Neural Mechanisms and Visual Behavior
Crustacean Experimental Systems in Neurobiology, 2002
Page 221. Visual Systems: Neural Mechanisms and Visual Behavior Raymon M. Glantz'and W. Jon.... more Page 221. Visual Systems: Neural Mechanisms and Visual Behavior Raymon M. Glantz'and W. Jon. P. Barnes 2 'Department of Biochemistry and Cell Biology, Rice University, Houston, Texas, USA 2Division of Environmental ...
Adhesion in Wet Environments: Frogs
Encyclopedia of Nanotechnology, 2012
SUMMARY 1. When a crayfish walks forward, its legs are not loaded equally. The third pair is the ... more SUMMARY 1. When a crayfish walks forward, its legs are not loaded equally. The third pair is the one exerting the largest vertically directed force, and the fourth legs produce the major part of the propulsive force. 2. Chronic recordings from a CSD2 nerve were made in crayfish walking on a treadmill. The activity of CSD2 rises during the stance phase. The activity peak at the beginning of the power stroke is especially pronounced in leg 4, which was studied in detail. The average spike frequency of CSD2 recordings during the first part of the power stroke is positively correlated with the initial increase in force produced by the leg. This suggests that CSD2 responds especially well to rapid load changes. 3. When a wire was fixed around a fourth leg such that it depressed the patch of compliant cuticle associated with CSD2, the activity of the sense organ was elevated throughout the entire step cycle, and the time between leg 4 touching the ground and leg 3 lifting off was more var...
Effects of light intensity and pattern contrast on the ability of the land crab, Cardisoma guanhumi, to separate optic flow-field components
Visual Neuroscience, 2004
Using a novel suite of computer-generated visual stimuli that mimicked components of optic flow, ... more Using a novel suite of computer-generated visual stimuli that mimicked components of optic flow, the visual responses of the tropical land crab, Cardisoma guanhumi, were investigated. We show that crabs are normally successful in distinguishing the rotational and translational components of the optic flow field, showing strong optokinetic responses to the former but not the latter. This ability was not dependant on the orientation of the crab, occurring both in “forwards-walking” and “sideways-walking” configurations. However, under conditions of low overall light intensity and/or low object/background contrast, the separation mechanism shows partial failure causing the crab to generate compensatory eye movements to translation, particularly in response to low-frequency (low-velocity) stimuli. Using this discovery, we then tested the ability of crabs to separate rotational and translational components in a combined rotation/translation flow field under different conditions. We demon...
Local mechanisms for the separation of optic flow-field components in the land crab, Cardisoma guanhumi: A role for motion parallax?
Visual Neuroscience, 2004
Although a number of global mechanisms have been proposed over the years that explain how crabs m... more Although a number of global mechanisms have been proposed over the years that explain how crabs might separate the rotational and translational components of their optic flow field, there has been no evidence to date that local mechanisms such as motion parallax are used in this separation. We describe here a study that takes advantage of a recently developed suite of computer-generated visual stimuli that creates a three-dimensional world surrounding the crab in which we can simulate translational and rotational optic flow. We show that, while motion parallax is not the only mechanism used in flow-field separation, it does play a role in the recognition of translational optic flow fields in that, under conditions of low overall light intensity and low contrast ratio when crabs find the distinction between rotation and translation harder, smaller eye movements occur in response to translation when motion parallax cues are present than when they are absent. Thus, motion parallax is o...
Journal of Experimental Biology, 2012
SUMMARY Tree frogs use adhesive toe pads for climbing on a variety of surfaces. They rely on wet ... more SUMMARY Tree frogs use adhesive toe pads for climbing on a variety of surfaces. They rely on wet adhesion, which is aided by the secretion of mucus. In nature, the pads will undoubtedly get contaminated regularly through usage, but appear to maintain their stickiness over time. Here, we show in two experiments that the toe pads of White's tree frogs (Litoria caerulea) quickly recover from contamination through a self-cleaning mechanism. We compared adhesive forces prior to and after contamination of (1) the whole animal on a rotatable platform and (2) individual toe pads in restrained frogs mimicking individual steps using a motorised stage. In both cases, the adhesive forces recovered after a few steps but this took significantly longer in single toe pad experiments from restrained frogs, showing that use of the pads increases recovery. We propose that both shear movements and a ‘flushing’ effect of the secreted mucus play an important role in shedding particles/contaminants.
Adhesion and allometry from metamorphosis to maturation in hylid tree frogs: a sticky problem
Journal of Zoology, 2006