Environmental Constraints on the Mechanics of Crawling and Burrowing Using Hydrostatic Skeletons

Dorgan, Kelly M.

doi:10.1007/s11340-010-9399-2

Cited by 15 publications

(14 citation statements)

References 55 publications

(74 reference statements)

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“…Means were not significantly different between experiments (P = 0.83). The mean burrowing speed of 1.6 mm s -1 measured in this study is nearly twice the 0.86 mm s -1 measured by Dorgan (2007). This difference is well outside the estimated 8.3% error (see "Validation and Parameterization") in the PIV measurement method and is likely to be an underestimate because only speed in the horizontal plane is measured here.…”

Section: Piv Experimentssupporting

confidence: 63%

“…I found no published measurements of burrowing speed for A. virens in sand. Dorgan (2007), however, measured an average burrowing speed of 0.86 mm s -1 for A. virens burrowing in gelatin as an analog for mud, providing a comparison for the burrowing speed measured in sand here. The distance of A. virens from the wall of its container was also measured as a test for results presented in an earlier study looking at differences in A. virens burrowing with distance from the wall in sand and mud (Du Clos et al 2013), as was the extent of sediment surface disturbed by burrowing.…”

mentioning

confidence: 60%

“…Second, gelatin has been used as a transparent surrogate for mud because of similarities in the material properties of the two media. Dorgan et al (2007) developed a linear-elastic fracture mechanics model of burrowing based on observations of Alitta virens burrowing in gelatin. Cryolite has been used as a transparent analogue for sand (Josephson and Flessa 1972;Francoeur and Dorgan 2014).…”

mentioning

confidence: 99%

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Visualizing subsurface burrowing by the polychaete Alitta virens with particle image velocimetry

Clos

2014

Limnology & Ocean Methods

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Infauna, animals that live in marine sediments, alter physical, chemical, and biological properties of their habitats through burrowing and other activities. Though important, infaunal behavior is notoriously difficult to study. Sediments are resistant to imaging techniques because they scatter radiation and are dense and opaque, and measurements made across aquarium walls are subject to artifacts. In this study, particle image velocimetry (PIV) was used to indirectly observe the activity of the marine polychaete Alitta virens as it burrowed in sand and to measure its horizontal burrowing speed. The mean burrowing speed of 1.6 mm s‐1 for A. virens is the first published measurement of subsurface burrowing speed in sand. The extent of surficial sediment displacement was also measured. This study demonstrates the utility of PIV as a method for observing burrowing behavior and suggests potential applications in laboratory and field studies of burrowing behavior.

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Section: Piv Experimentssupporting

confidence: 63%

mentioning

confidence: 60%

mentioning

confidence: 99%

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Visualizing subsurface burrowing by the polychaete Alitta virens with particle image velocimetry

Clos

2014

Limnology & Ocean Methods

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“…The biomechanics of soil penetration by earthworms relies on peristaltic synchronization of contracting and stretching muscle fibers (Quillin, 2000) that constitute the earthworm's hydrostatic skeleton (Dorgan, 2010). Cyclic penetration–expansion steps are driven by peristalsis of the earthworm's pressurized colonic fluid (Dorgan et al, 2007; Murphy and Dorgan, 2011).…”

Section: Methodsmentioning

confidence: 99%

Mechanics and Energetics of Soil Penetration by Earthworms and Plant Roots: Higher Rates Cost More

Ruiz

Schymanski

2017

Vadose Zone Journal

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Core Ideas We quantified mechanical differences between earthworms and roots using penetration rates. Mechanical modeling was justified by direct imaging of both plant roots and earthworms. We validated model predictions with rate‐controlled miniature cone penetrometer experiments. Earthworm burrows are dominantly formed through mechanical processes, not ingestion. We outline mechanical and energetic limitations for a range of water contents. We quantified the mechanics and energetics of soil penetration by burrowing earthworms and growing plant roots considering different penetration rates and soil mechanical properties. The mechanical model considers cavity expansion by cone‐like penetration into a viscoelastic soil material in which penetration rates affect the resulting forces and hence the mechanical energy required. To test the predicted penetration rate effects on forces and energetics, we conducted rate‐controlled cone penetration experiments across rates ranging from 1 to 200 μm s−1 to determine the mechanical resistance forces for cone geometries similar to plant roots and earthworms. These measurements also enabled inverse estimation of soil rheological parameters that were in good agreement with literature values for similar soils and water contents. The results suggest that higher soil penetration rates typical for earthworm activity (about 200 μm s−1) may significantly increase resistance forces and energy expenditure by up to threefold relative to slower penetration rates of plant roots (0.2 μm s−1) for similar soil properties and geometries. Another important mechanical difference between earthworms and roots is the radial pressures that earthworms' hydro‐skeleton exerts (<230 kPa), whereas plant roots may exert radial pressures exceeding 1 MPa. These inherent differences in burrowing rates and expansion pressures may significantly extend the range of conditions suitable for root growth in drier and compacted soil compared to earthworm activity. Results suggest that the mechanical energy costs of soil bioturbation under agricultural intensification and drier climate could greatly increase the energetic costs of these ecologically important soil structure‐forming bioprocesses.

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“…Crawling with a hydrostatic skeleton has been found by several authors to be costly (Denny, 1980;Casey, 1991;Berrigan and Lighton, 1993). Generalizing locomotory strategies across animals with hydrostatic skeletons is questionable, however, as the mechanics of burrowing in saturated sediments differ substantially from those of terrestrial crawling (Dorgan, 2010).…”

Section: Introductionmentioning

confidence: 99%

Energetics of burrowing by the cirratulid polychaeteCirriformia moorei

Dorgan

Lefebvre

Stillman

et al. 2011

Journal of Experimental Biology

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SUMMARYBurrowing through marine sediments has been considered to be much more energetically expensive than other forms of locomotion, but previous studies were based solely on external work calculations and lacked an understanding of the mechanical responses of sediments to forces applied by burrowers. Muddy sediments are elastic solids through which worms extend crackshaped burrows by fracture. Here we present data on energetics of burrowing by Cirriformia moorei. We calculated the external energy per distance traveled from the sum of the work to extend the burrow by fracture and the elastic work done to displace sediment as a worm moves into the newly formed burrow to be 9.7Jkg -1 m -1 in gelatin and 64Jkg -1 m -1 in sediment, much higher than for running or walking. However, because burrowing worms travel at slow speeds, the increase in metabolic rate due to burrowing is predicted to be small. We tested this prediction by measuring aerobic metabolism (oxygen consumption rates) and anaerobic metabolism (concentrations of the anaerobic metabolite tauropine and the energy-storage molecule phosphocreatine) of C. moorei. None of these components was significantly different between burrowing and resting worms, and the low increases in oxygen consumption rates or tauropine concentrations predicted from external work calculations were within the variability observed across individuals. This result suggests that the energy to burrow, which could come from aerobic or anaerobic sources, is not a substantial component of the total metabolic energy of a worm. Burrowing incurs a low cost per unit of time.Supplementary material available online at

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Environmental Constraints on the Mechanics of Crawling and Burrowing Using Hydrostatic Skeletons

Cited by 15 publications

References 55 publications

Visualizing subsurface burrowing by the polychaete Alitta virens with particle image velocimetry

Visualizing subsurface burrowing by the polychaete Alitta virens with particle image velocimetry

Mechanics and Energetics of Soil Penetration by Earthworms and Plant Roots: Higher Rates Cost More

Energetics of burrowing by the cirratulid polychaeteCirriformia moorei

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