Extreme positive allometry of animal adhesive pads and the size limits of adhesion-based climbing
- aDepartment of Zoology, University of Cambridge, Cambridge CB2 3EJ, United Kingdom;
- bSchool of Science and Engineering, University of the Sunshine Coast, Sippy Downs, QLD 4556, Australia;
- cDepartment of Life Sciences, Anglia Ruskin University, Cambridge CB1 1PT, United Kingdom;
- dBiology Department, University of Massachusetts Amherst, Amherst, MA 01003;
- eDepartment of Polymer Science and Engineering, University of Massachusetts Amherst, Amherst, MA 01003
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Edited by David B. Wake, University of California, Berkeley, CA, and approved December 11, 2015 (received for review October 7, 2015)

Significance
How adhesive forces can be scaled up from microscopic to macroscopic levels is a central problem for biological and bio-inspired adhesives. Here, we elucidate how animals with sticky footpads cope with large body sizes. We find an extreme positive allometry of footpad area across all 225 species studied, implying a 200-fold increase of relative pad area from mites to geckos. Within groups, however, pads were almost isometric, but their adhesive strength increased with size, inconsistent with existing models. Extrapolating the observed scaling, we show that to support a human’s body weight, an unrealistic 40% of the body surface would have to be covered with adhesive pads, suggesting that anatomical constraints may prohibit the evolution of adhesion-based climbers larger than geckos.
Abstract
Organismal functions are size-dependent whenever body surfaces supply body volumes. Larger organisms can develop strongly folded internal surfaces for enhanced diffusion, but in many cases areas cannot be folded so that their enlargement is constrained by anatomy, presenting a problem for larger animals. Here, we study the allometry of adhesive pad area in 225 climbing animal species, covering more than seven orders of magnitude in weight. Across all taxa, adhesive pad area showed extreme positive allometry and scaled with weight, implying a 200-fold increase of relative pad area from mites to geckos. However, allometric scaling coefficients for pad area systematically decreased with taxonomic level and were close to isometry when evolutionary history was accounted for, indicating that the substantial anatomical changes required to achieve this increase in relative pad area are limited by phylogenetic constraints. Using a comparative phylogenetic approach, we found that the departure from isometry is almost exclusively caused by large differences in size-corrected pad area between arthropods and vertebrates. To mitigate the expected decrease of weight-specific adhesion within closely related taxa where pad area scaled close to isometry, data for several taxa suggest that the pads’ adhesive strength increased for larger animals. The combination of adjustments in relative pad area for distantly related taxa and changes in adhesive strength for closely related groups helps explain how climbing with adhesive pads has evolved in animals varying over seven orders of magnitude in body weight. Our results illustrate the size limits of adhesion-based climbing, with profound implications for large-scale bio-inspired adhesives.
Footnotes
- ↵1To whom correspondence should be addressed. Email: dl416{at}cam.ac.uk.
Author contributions: D.L. and W.F. designed research; D.L., C.J.C., A.D., and C.-Y.K. performed research; D.L. and C.J.C. analyzed data; and D.L., C.J.C., A.J.C., D.J.I., and W.F. wrote the paper.
The authors declare no conflict of interest.
This article is a PNAS Direct Submission.
This article contains supporting information online at www.pnas.org/lookup/suppl/doi:10.1073/pnas.1519459113/-/DCSupplemental.
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