Eye-specific learning of routes and “signposts” by walking honeybees

This study investigates the honeybee's ability to learn routes based on visual stimuli presented to a single eye, and to then navigate these routes using the other (naive) eye. Bees were trained to walk through a narrow tunnel carrying visual stimuli on the two walls. At the end of the tunnel the bees had to choose between two arms, one of which led to a feeder. In a first experiment, bees had to learn to choose the left arm to get a reward when the right wall carried a yellow grating, but the right arm when the left wall carried a blue grating. The bees learned this task well, indicating that stimuli encountered by different eyes could be associated with different routes. In a second experiment, bees had to turn left when the right eye saw a blue grating, but to the right when the same eye saw a yellow grating. They also learned this task well. In subsequent tests, they chose the correct arm even when these gratings were presented to the untrained eye. These results suggest that there is interocular transfer of route-specific learning with respect to visual stimuli that function as navigational “signposts”. Accepted: 18 December 1997     

Movement behavior of high-heeled walking: how does the nervous system control the ankle joint during an unstable walking condition?

The human locomotor system is flexible and enables humans to move without falling even under less than optimal conditions. Walking with high-heeled shoes constitutes an unstable condition and here we ask how the nervous system controls the ankle joint in this situation? We investigated the movement behavior of high-heeled and barefooted walking in eleven female subjects. The movement variability was quantified by calculation of approximate entropy (ApEn) in the ankle joint angle and the standard deviation (SD) of the stride time intervals. Electromyography (EMG) of the soleus (SO) and tibialis anterior (TA) muscles and the soleus Hoffmann (H-) reflex were measured at 4.0 km/h on a motor driven treadmill to reveal the underlying motor strategies in each walking condition. The ApEn of the ankle joint angle was significantly higher (p<0.01) during high-heeled (0.38±0.08) than during barefooted walking (0.28±0.07). During high-heeled walking, coactivation between the SO and TA muscles increased towards heel strike and the H-reflex was significantly increased in terminal swing by 40% (p<0.01). These observations show that high-heeled walking is characterized by a more complex and less predictable pattern than barefooted walking. Increased coactivation about the ankle joint together with increased excitability of the SO H-reflex in terminal swing phase indicates that the motor strategy was changed during high-heeled walking. Although, the participants were young, healthy and accustomed to high-heeled walking the results demonstrate that that walking on high-heels needs to be controlled differently from barefooted walking. We suggest that the higher variability reflects an adjusted neural strategy of the nervous system to control the ankle joint during high-heeled walking.     

Did knuckle walking evolve twice?

Although African great apes share a similar quadrupedal locomotor behaviour, there are marked differences in hand morphology and size between the species. Hence, whilst all three species (two genera) of African ape frequently knuckle walk as adults, debate remains as to whether this behaviour is derived from a common ancestor or whether it evolved in parallel in chimpanzees and gorillas. This exploratory morphometric study of the sub-adult and adult wrist of these two genera aims to contribute to this debate. A total of twenty-seven dimensions of the lunate, triquetral, hamate and capitate of sub-adult and adult Pan troglodytes and Gorilla gorilla were analysed in order to determine whether carpal dimensions are generally ontogenetically scaled, and whether differences in growth trajectories, or length of growth, and adult morphologies can be explained by behavioural differences between the two species. Only 56% of all dimensions studied were ontogenetically scaled in sub-adults and some of these dimensions exhibit differing adult proportions between the two species. In general, the dimensions analysed fell into two categories: Pan and Gorilla either follow the same growth trajectories (Pattern A) or the Pan reduced major axis (RMA) regressions were significantly transposed above those of Gorilla (Pattern B). Additionally, it was found that Gorilla carpals appear to cease growing relatively earlier than those of Pan. While a small number of differences, notably those of the lunate, can be accounted for by differences in behaviour between the species, the majority of differences indicate heterochronic modifications of development during evolution, which correspond to kinematic differences in knuckle walking between the African great apes. In light of morphological, behavioural and ecological data currently available it is parsimonious to suggest that knuckle walking has evolved in parallel in the two lineages.     

Is slow walking more stable?

Several efforts have been made to study gait stability using measures derived from nonlinear time-series analysis. The maximum finite time Lyapunov exponent (λ_max) quantifies how a system responds to an infinitesimally small perturbation. Recent studies suggested that slow walking leads to lower λ_max values, and thus is more stable than fast walking, but these studies suffer from methodological limitations. We studied the effects of walking speed on the amount of kinematic variability and stability in human walking. Trunk motions of 15 healthy volunteers were recorded in 3D during 2 min of treadmill walking at different speeds. From those time series, maximum Lyapunov exponents, indicating short-term and long-term divergence (λ_S-stride and λ_L-stride), and mean standard deviation (MeanSD) were calculated. λ_S-stride showed a linear decrease with increasing speed for forward–backward (AP) movements and quadratic effects (inverted U-shaped) for medio-lateral (ML) and up–down (VT) movements. λ_L-stride showed a quadratic effect (inverted U-shaped) of walking speed for AP movements, a linear decrease for ML movements, and a linear increase for VT movements. Moreover, positive correlations between λ_S and MeanSD were found for all directions, while λ_L-stride and MeanSD were correlated negatively in the AP direction. The different effects of walking speed on λ_S-stride and λ_L-stride for the different planes suggest that slow walking is not necessarily more stable than fast walking. The absence of a consistent pattern of correlations between λ_L-stride and MeanSD over the three directions suggests that variability and stability reflect, at least to a degree, different properties of the dynamics of walking.     

Atomistic probing of aptameric binding of CD19 outer membrane domain reveals an “aptamer walking” mechanism

We propose an integrated structural approach to search potential aptamer molecules for targeting cancer receptor proteins. We used the outer cellular domain of the B-lymphocyte antigen, CD19, as the target for this study. First, using available protein-aptamer structures deposited in the protein data bank as resources, structural annotation was performed to seek the most probable binding aptamer and its potential initial configuration to the CD19 structure. Using this initial structure, molecular dynamics (MD) simulations were performed for adjustment of the aptamer-binding. During this process, we observed an “aptamer walking” mechanism of the binding of the single-stranded RNA-aptamer to CD19: the aptamer molecule gradually adjusts its configurations and shifts toward favorable binding positions. However, the target molecule CD19 maintained a relatively stable conformation during this process. The interface area between the RNA-aptamer and CD19 increased from less than 8 nm² to over 12 nm² during a 2-μs MD simulation. Using a stable binding pose as the starting structure, we manually mutated the RNA-aptamer to a DNA-aptamer and found that the interface area was further increased to over 16 nm², indicating a stronger affinity compared to the RNA-aptamer. The RNA- and DNA-aptamers and their stable binding-poses to the CD19 molecule may be used as templates in designing potential aptamer molecules that target the B-cell marker molecule CD19 with enhanced specificity and stability.     

Tension receptors on the apodemes of muscles in the walking legs of the crab,Cancer magister †

1. Mechanoreceptors monitoring tension in working muscles are described in the Decapoda Crustacea.

2. The receptors are associated with apodemes of muscles in the walking leg and are well‐developed in the extensor and flexor of the meropodite (Figures 1, 2).

3. The unbranched dendrites of the receptor neurones innervate the tissues surrounding the insertions of the muscle fibres (Figures 3, 4, 5(A)).

4. The receptors show spontaneous activity with the M‐C joint at resting position and this activity increases when the muscle is stretched by holding the joint at a different position (Figure 7).

5. Isometric tension increase in the muscle recruits sensory units (Figures 8, 10(A)) and increases the activity of units firing (Figure 9).

6. Apodeme receptors may be an entirely distinct input channel from chordotonal organs (Figure 10(B,C)). Joint movements produced by a standard muscle stimulus against increasing loads reveal very different responses (Figure 11).

7. Attempts to determine whether chordotonal organs (CP1, Figures 5(B), 6) monitor isometric muscle tension (Figure 12) suggest possible complexities in their dynamic responses.

8. Abbreviations used in this paper are FASN flexor apodeme sensory nerve, EASN extensor apodeme sensory nerve, BASN bender apodeme sensory nerve, and OASN opener apodeme sensory nerve.     

Proprioception, gait kinematics, and rate of loading during walking: are they related?

The cyclic nature of walking can lead to repetitive stress and associated complications due to the rate of loading (ROL) experienced by the body at the initial contact of the foot with the ground. An individual's gait kinematics at initial contact has been suggested to give rise to the ROL, and a repetitive, high ROL may lead to several disorders, including osteoarthritis. Additionally, proprioception, the feedback signaling of limb position and movement, may play a role in how the foot strikes the ground and thus, the ROL. Our goal was to explore the relationship between proprioception, gait kinematics and ROL. Thirty-eight women were recruited for gait analysis, and the gait characteristics 50 ms prior to and at initial contact were examined. Two proprioception tests, joint angle reproduction and threshold to detect passive motion were used to examine the subject's proprioceptive acuity. Our results indicate that individuals with a larger knee angle (i.e., greater extension) 50 ms prior to initial contact (IC) experience a higher ROL during gait and have poorer proprioceptive scores. However, it remains unclear whether poor proprioception causes a high ROL or if a high ROL damages the mechanoreceptors involved in proprioception, but the apparent relationship is significant and warrants further investigation.     

Do patients with diabetic neuropathy use a higher proportion of their maximum strength when walking?

Diabetic patients have an altered gait strategy during walking and are known to be at high risk of falling, especially when diabetic peripheral neuropathy is present. This study investigated alterations to lower limb joint torques during walking and related these torques to maximum strength in an attempt to elucidate why diabetic patients are more likely to fall. 20 diabetic patients with moderate/severe peripheral neuropathy (DPN), 33 diabetic patients without peripheral neuropathy (DM), and 27 non-diabetic controls (Ctrl) underwent gait analysis using a motion analysis system and force plates to measure kinetic parameters. Lower limb peak joint torques and joint work done (energy expenditure) were calculated during walking. The ratio of peak joint torques and individual maximum joint strengths (measured on a dynamometer) was then calculated for 59 of the 80 participants to yield the ‘operating strength’ for those participants. During walking DM and DPN patients showed significantly reduced peak torques at the ankle and knee. Maximum joint strengths at the knee were significantly less in both DM and DPN groups than Ctrls, and for the DPN group at the ankle. Operating strengths were significantly higher at the ankle in the DPN group compared to the Ctrls. These findings show that diabetic patients walk with reduced lower limb joint torques; however due to a decrement in their maximum ability at the ankle and knee, their operating strengths are higher. This allows less reserve strength if responding to a perturbation in balance, potentially increasing their risk of falling.     

Kinesin: walking, crawling or sliding along?

Kinesins are microtubule-based motor proteins that are involved in cargo transport and mitosis. They are called "motors" because they convert chemical energy to mechanical energy (i.e. force and motion). They use the energy of ATP hydrolysis for their enzymatic processes by walking on microtubules. However, the mechanism underlying their motion has been unclear. Recently, conventional kinesin, which was the first-identified member of the family, has been shown to walk by swapping its two heads in a "hand-over-hand" mechanism. There is also experimental evidence supporting an asymmetric walking of kinesin in which two identical heads of the motor take alternate slow and fast steps. Other cargo-carrier and mitotic kinesins remain uninvestigated and are of great interest to biophysicists.     

Prothoracic DUM neurons of the cricket Gryllus bimaculatus — responses to natural stimuli and activity in walking behavior

Summary Responses to sensory stimuli and spike activity uring walking were investigated in bilaterally symmetrical dorsal unpaired median (DUM) neurons of the cricket. Intracellular recordings within the prothoracic ganglion were made either in restrained animals or in stationary walking specimens whilst parameters of their intended locomotion were measured. Three types of DUM cells were distinguished morphologically and physiologically. DUMa neurons send axons through segmental nerves. They often generated spontaneously large action potentials with low frequencies. Most DUMa neurons showed multimodal sensitivity, preferentially to cereal wind puffs and 15 kHz sound. Mean latencies ranged from 25 to 349 ms. Their large intraindividual variability could be correlated with behavioral modes during walking. Generally, the spike frequency increased with increased forward speed, while it was not related to turning. DUMb neurons projected either through the anterior or posterior connectives, but seemed physiologically similar to DUMa neurons. DUMc neurons were H-shaped with axons in both pairs of connectives. No external stimulus led to discrete spikes, but the regular spontaneous activity was modulated following cereal wind puffs to a restrained animal. During wind evoked escape the spike activity of another DUMc cell was modulated in phase with the rhythmic running behavior. The possibly different functions of DUMa and DUMc neurons during walking are discussed.     

Dead land walking: the value of continued conservation efforts in South Florida’s imperiled pine rocklands

Pine rocklands are deeply imperiled habitats restricted to South Florida and the Caribbean. In South Florida, more than 98% of pine rockland habitat has been destroyed in the past century (outside of Everglades National Park). Due to their proximity to human populations, management options in the remaining fragments are sometimes limited, and fires that are necessary to maintain healthy habitat structure are often excluded. Despite these pressures, conservation initiatives in pine rocklands have been surprisingly successful, and plant extinction has been avoided. In the coming decades, however, sea-level rise threatens to all but eliminate the pine rocklands, and efforts to preserve their many endemic species will likely fail. We synthesize the results of numerous ecological studies and review the successes and failures of conservation in South Florida’s pine rocklands. Further, we illustrate the value of continued conservation efforts, and provide direction in the light of the habitats long-term fate. We advocate the increased use of prescribed fire and, as the effects of climate change become more apparent, the translocation of some endemic species. Finally, we acclaim pine rocklands as a model system for studying how plant communities respond to environmental change. South Florida’s fragmented landscape, with shifting gradients of elevation, salinity, inundation and nutrient availability, should continue to inspire ecologists to address important questions, and better prepare the region, and the world, for the challenges of the coming decades.     

Does adiposity affect muscle function during walking in children?

The biomechanical mechanisms responsible for the altered gait in obese children are not well understood, particularly as they relate to increases in adipose tissue. The purpose of this study was to test the hypotheses that as body-fat percentage (BF%) increased: (1) knee flexion during stance would decrease while pelvic obliquity would increase; (2) peak muscle forces normalized to lean-weight would increase for gluteus medius, gastrocnemius, and soleus, but decrease for the vasti; and (3) the individual muscle contributions to center of mass (COM) acceleration in the direction of their primary function(s) would not change for gluteus medius, gastrocnemius, and soleus, but decrease for the vasti. We scaled a musculoskeletal model to the anthropometrics of each participant (n=14, 8–12 years old, BF%: 16–41%) and estimated individual muscle forces and their contributions to COM acceleration. BF% was correlated with average knee flexion angle during stance (r=−0.54, p=0.024) and pelvic obliquity range of motion (r=0.78, p<0.001), as well as with relative vasti (r=−0.60, p=0.023), gluteus medius (r=0.65, p=0.012) and soleus (r=0.59, p=0.026) force production. Contributions to COM acceleration from the vasti were negatively correlated to BF% (vertical— r=−0.75, p=0.002, posterior— r=−0.68, p=0.008), but there were no correlation between BF% and COM accelerations produced by the gastrocnemius, soleus and gluteus medius. Therefore, we accept our first, partially accept our second, and accept our third hypotheses. The functional demands and relative force requirements of the hip abductors during walking in pediatric obesity may contribute to altered gait kinematics.     

Can treadmill walking be used to assess propulsion generation?

Instrumented treadmills offer significant advantages for analysis of human locomotion, including recording consecutive steady-state gait cycles, precisely controlling walking speed, and avoiding force plate targeting. However, some studies of hemiparetic walking on a treadmill have suggested that the moving treadmill belt may fundamentally alter propulsion mechanics. Any differences in propulsion mechanics during treadmill walking would be problematic since recent studies assessing propulsion have provided fundamental insight into hemiparetic walking. The purpose of this study was to test the hypothesis that there would be no difference in the generation of anterior/posterior (A/P) propulsion by performing a carefully controlled comparison of the A/P ground reaction forces (GRFs) and impulses in healthy adults during treadmill and overground walking. Gait data were collected from eight subjects walking overground and on a treadmill with speed and cadence controlled. Peak negative and positive horizontal GRFs in early and late stance, respectively, were reduced by less than 5% of body weight (p<0.05) during treadmill walking compared to overground walking. The magnitude of the braking impulse was similarly lower (p<0.05) during treadmill walking, but no significant difference was found between propulsion impulses. While there were some subtle differences in A/P GRFs between overground and treadmill walking, these results suggest there is no fundamental difference in propulsion mechanics. We conclude that treadmill walking can be used to investigate propulsion generation in healthy and by implication clinical populations.     

Relationships between unitary motor and sensory activity in the walking legs of cancer pagurus (Decapoda,Brachyura)†

Muscular tension and sensory activity in the flexor apodeme sensory nerve were recorded during stimulation of single motor afferents innervating the M‐C flexor. Muscular tension and unitary sensory activity both varied, depending upon the motor fiber stimulated. Differences in the abililty of individual motor fibers to elicit sensory activity were only partially accounted for by differences in tension development. Some tension afferent units were more readily excited by a muscular contraction elicited by one motor axon than they were by another, even when the tension elicited by the more effective motor fiber was less than that evoked by the less effective efferent.     

Does cypermethrin affect enzyme activity,respiration rate and walking behavior of the maize weevil (Sitophilus zeamais)?

Insecticides cause a range of sub‐lethal effects on targeted insects, which are frequently detrimental to them. However, targeted insects are able to cope with insecticides within sub‐lethal ranges, which vary with their susceptibility. Here we assessed the response of three strains of the maize weevil Sitophilus zeamais Motschulsky (Coleoptera: Curculionidae) to sub‐lethal exposure to the pyrethoid insecticide cypermethrin. We expected enzyme induction associated with cypermethrin resistance since it would aid the resistant insects in surviving such exposure. Lower respiration rate and lower activity were also expected in insecticide‐resistant insects since these traits are also likely to favor survivorship under insecticide exposure. Curiously though, cypermethrin did not affect activity of digestive and energy metabolism enzymes, and even reduced the activity of some enzymes (particularly for cellulase and cysteine‐proteinase activity in this case). There was strain variation in response, which may be (partially) related to insecticide resistance in some strains. Sub‐lethal exposure to cypermethrin depressed proteolytic and mainly cellulolytic activity in the exposed insects, which is likely to impair their fitness. However, such exposure did not affect respiration rate and walking behavior of the insects (except for the susceptible strain where walking activity was reduced). Walking activity varies with strain and may minimize insecticide exposure, which should be a concern, particularly if associated with (physiological) insecticide resistance.     

Development and validation of a new method to measure walking speed in free-living environments using the actibelt® platform

Walking speed is a fundamental indicator for human well-being. In a clinical setting, walking speed is typically measured by means of walking tests using different protocols. However, walking speed obtained in this way is unlikely to be representative of the conditions in a free-living environment. Recently, mobile accelerometry has opened up the possibility to extract walking speed from long-time observations in free-living individuals, but the validity of these measurements needs to be determined. In this investigation, we have developed algorithms for walking speed prediction based on 3D accelerometry data (actibelt?) and created a framework using a standardized data set with gold standard annotations to facilitate the validation and comparison of these algorithms. For this purpose 17 healthy subjects operated a newly developed mobile gold standard while walking/running on an indoor track. Subsequently, the validity of 12 candidate algorithms for walking speed prediction ranging from well-known simple approaches like combining step length with frequency to more sophisticated algorithms such as linear and non-linear models was assessed using statistical measures. As a result, a novel algorithm employing support vector regression was found to perform best with a concordance correlation coefficient of 0.93 (95%CI 0.92-0.94) and a coverage probability CP1 of 0.46 (95%CI 0.12-0.70) for a deviation of 0.1 m/s (CP2 0.78, CP3 0.94) when compared to the mobile gold standard while walking indoors. A smaller outdoor experiment confirmed those results with even better coverage probability. We conclude that walking speed thus obtained has the potential to help establish walking speed in free-living environments as a patient-oriented outcome measure.     

The influence of a 1-year programme of brisk walking on endurance fitness and body composition in previously sedentary men aged 42–59 years

This study examined the influence of a 1-year brisk walking programme on endurance fitness and the amount and distribution of body fat in a group of formerly sedentary men. Seventy-two males, aged 42–59 years, body mass index 25.2 (0.3) kg·m^–2 [mean (SEM)], were randomly allocated to either a walking group (n = 48) or control group (n = 24). Brisk walking speed was evaluated using a 1.6-km track walk. Changes in endurance fitness were assessed by measuring blood lactate concentration and heart rate during submaximal treadmill walking. Body composition was determined by hydrostatic weighing and anthropometry; energy intake was assessed from 7-day weighed food inventories. Differences in the response of walkers and controls were examined using two-way analyses of variance. Forty-two walkers (87.5%) completed the study and averaged 27.9 (1.4) min·day^–1 of brisk walking (range 11–46). Brisk walking speed averaged 1.95 (0.03) m·s^–1 and elicited approximately 68 (1) % of maximum heart rate. Heart rate and blood lactate concentration during submaximal treadmill walking were significantly reduced in the walkers after 3, 6 and 12 months and the oxygen uptake at a reference blood lactate concentration of 2 mmol·l^–1 was increased by 3.2 ml·kg^–1. min^–1 (14.9%) in the walkers at 6 months (P< 0.01). Although skinfold thicknesses at anterior thigh and medial calf sites decreased significantly for the walkers, the response of the two groups did not differ significantly for other body composition variables or for energy intake.