Many ecological factors influence animal movement, including properties of the media that they move on or through. Animals moving in terrestrial environments encounter conditions that can be challenging for generating propulsion and maintaining stability, such as inclines and deformable substrates that can cause slipping and sinking. In response, tetrapods tend to adopt a more crouched posture and lower their center of mass on inclines and increase the surface area of contact on deformable substrates, such as sand. Many amphibious fishes encounter the same challenges when moving on land, but how these finned animals modulate their locomotion with respect to different environmental conditions and how these modifications compare with those seen within tetrapods is relatively understudied. Mudskippers (Gobiidae: Oxudercinae) are a particularly noteworthy group of amphibious fishes in this context given that they navigate a wide range of environmental conditions, from flat mud to inclined mangrove trees. They use a unique form of terrestrial locomotion called “crutching,” where their pectoral fins synchronously lift and vault the front half of the body forward before landing on their pelvic fins, while the lower half of the body and tail are kept straight. However, recent work has shown that mudskippers modify some aspects of their locomotion when crutching on deformable surfaces, particularly those at an incline. For example, on inclined dry sand, mudskippers bent their bodies laterally and curled and extended their tails to potentially act as a secondary propulsor and/or anti-slip device. In order to gain a more comprehensive understanding of the functional diversity and context-dependency of mudskipper crutching, we compared their kinematics on different combinations of substrate types (solid, mud, and dry sand) and inclines (0°, 10°, and 20°). In addition to increasing lateral bending on deformable and inclined substrates, we found that mudskippers increased the relative contact time and contact area of their paired fins, while becoming more crouched, which are responses comparable to those seen in tetrapods and other amphibious fish. Mudskippers on these substrates also exhibited previously undocumented behaviors, such as extending and adpressing the distal portions of their pectoral fins more anteriorly, dorsoventrally bending their trunk, “belly-flopping” on sand, and “gripping” the mud substrate with their pectoral fin rays. Our study highlights potential compensatory mechanisms shared among vertebrates in terrestrial environments while also illustrating that locomotor flexibility and even novelty can emerge when animals are challenged with environmental variation.
The spring-mass model is a model of locomotion aimed at giving the essential mathematical laws of the trajectory of the center of mass of an animal during bouncing gaits, such as hopping (one-dimensional) and running (two-dimensional). This reductionist mechanical system has been extensively investigated for locomotion over horizontal surfaces, whereas it has been largely neglected on other ecologically relevant surfaces, including inclines. For example, how the degree of inclination impacts the dynamics of the center of mass of the spring-mass model has not been investigated thoroughly. In this work, we derive a mathematical model which extends the spring-mass model to inclined surfaces. Among our results, we derive an approximate solution of the system, assuming a small angular sweep of the limb and a small spring compression during stance, and show that this approximation is very accurate, especially for small inclinations of the ground. Furthermore, we derive theoretical bounds on the difference between the Lagrangian and Lagrange equations of the true and approximate systems, and discuss locomotor stability questions of the approximate solutions. We test our models through a sensitivity analysis using parameters relevant to the locomotion of bipedal animals (quail, pheasant, guinea fowl, turkey, ostrich, and humans) and compare our approximate solution to the numerically derived solution of the exact system. We compare the two-dimensional spring-mass model on inclines with the one-dimensional spring-mass model to which it reduces under the limit of no horizontal velocity; we compare the two-dimensional spring-mass model on inclines with the inverted pendulum model on inclines towards which it converges in the case of high stiffness-to-mass ratio. We include comparisons with historically prevalent no-gravity approximations of these models, as well. The insights we have gleaned through all these comparisons and the ability of our approximation to replicate some of the kinematic changes observed in animals moving on different inclines (e.g., reduction in vertical oscillation of the center of mass and decreased stride length) underline the valuable and reasonable contributions that very simple, reductionist models, like the spring-mass model, can provide.
more » « less- NSF-PAR ID:
- 10367949
- Publisher / Repository:
- Oxford University Press
- Date Published:
- Journal Name:
- Integrative and Comparative Biology
- ISSN:
- 1540-7063
- Format(s):
- Medium: X
- Sponsoring Org:
- National Science Foundation
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Key points Vision plays a crucial role in guiding locomotion in complex environments, but the coordination between gaze and stride is not well understood.
The coordination of gaze shifts, fixations, constant gaze and slow gaze with strides in cats walking on different surfaces were examined.
It was found that gaze behaviours are coordinated with strides even when walking on a flat surface in the complete darkness, occurring in a sequential order during different phases of the stride.
During walking on complex surfaces, gaze behaviours are typically more tightly coordinated with strides, particularly at faster speeds, only slightly shifting in phase.
These findings indicate that the coordination of gaze behaviours with strides is not vision‐driven, but is a part of the whole body locomotion synergy; the visual environment and locomotor task modulate it. The results may be relevant to developing diagnostic tools and rehabilitation approaches for patients with locomotor deficits.
Abstract Vision plays a crucial role in guiding locomotion in complex environments. However, the coordination between the gaze and stride is not well understood. We investigated this coordination in cats walking on a flat surface in darkness or light, along a horizontal ladder and on a pathway with small stones. We recorded vertical and horizontal eye movements and 3‐D head movement, and calculated where gaze intersected the walkway. The coordination of gaze shifts away from the animal, gaze shifts toward, fixations, constant gaze, and slow gaze with strides was investigated. We found that even during walking on the flat surface in the darkness, all gaze behaviours were coordinated with strides. Gaze shifts and slow gaze toward started in the beginning of each forelimb's swing and ended in its second half. Fixations peaked throughout the beginning and middle of swing. Gaze shifts away began throughout the second half of swing of each forelimb and ended when both forelimbs were in stance. Constant gaze and slow gaze away occurred in the beginning of stance. However, not every behaviour occurred during every stride. Light had a small effect. The ladder and stones typically increased the coordination and caused gaze behaviours to occur 3% earlier in the cycle. At faster speeds, the coordination was often tighter and some gaze behaviours occurred 2–16% later in the cycle. The findings indicate that the coordination of gaze with strides is not vision‐driven, but is a part of the whole body locomotion synergy; the visual environment and locomotor task modulate it.
