The coordination of force oscillations and of leg movement in a walking insect (Carausius morosus)

As in the preceding paper stick insects walk on a treadwheel and different legs are put on platforms fixed relative to the insect's body. The movement of the walking legs is recorded in addition to the force oscillations of the standing legs. The coordination between the different legs depends upon the number and arrangement of the walking legs and the legs standing on platforms. In most experimental situations one finds a coordination which is different from that of a normal walking animal.Supported by DFG (Cr 58/1)     

The contributions of diverse sense organs to the control of leg movement by a walking insect

Summary During locomotion, stick insectsCarausius morosus, place the tarsus of the rear leg near the tarsus of the ipsilateral middle leg, whatever the position of the latter. This adjustment by the hind leg requires that it receive information on the actual position of the middle leg tarsus. It is shown by ablation experiments that such information is contributed by the following proprioceptors of the middle leg: the ventral and dorsal coxal hairplates, the coxal hair rows, the trochanteral hairplate and the femoral chordotonal organ. Additional information comes from other, as yet unidentified, sense organs. Several alternatives are considered to explain how the signals from the diverse sense organs of the subcoxal joint might be combined in computing the target position for the protracting hind leg. The experimental results support the hypothesis that the signals are added nonlinearly and that a signal deviating from the majority pattern is weighted less.Abbreviations cxHPu ventral coxal hairplate - cxHPd dorsal coxal hairplate - trHP trochanteral hairplate - HR hair row - feCO femoral chordotonal organ - AEP anterior extreme position     

The central connections and actions during walking of tibial campaniform sensilla in the locust

Strain acting on the exoskeleton of insects is monitored by campaniform sensilla. On the tibia of a mesothoracic leg of the locust (Schistocerca gregaria) there are three groups of campaniform sensilla on the proximo-dorsal surface. This study analyses the responses of the afferents from one group, their connections with central neurones and their actions during walking.The afferents of the campaniform sensilla make direct excitatory connections with flexor tibiae motor neurones. They also make direct connections with particular spiking local interneurones that make direct inhibitory output connections with the slow extensor tibiae motor neurone.During walking extension movements of the tibiae during stance produce longitudinal tensile forces on the dorsal tibia that peak during mid stance before returning to zero prior to swing. This decline in tension can activate the campaniform sensilla. In turn this would lead to an inhibition of the extensor tibiae motor neurone and an excitation of the flexor tibiae motor neurones. This, therefore, aids the transition from stance to swing. During turning movements, the tibia is flexed and the dorsal surface is put under compression. This can also activate some of campaniform sensilla whose effect on the flexor motor neurones will reinforce the flexion of the tibia.     

Possible functional roles of phase resetting during walking

The walking rhythm is known to show phase shift or "reset" in response to external impulsive perturbations. We tried to elucidate functional roles of the phase reset possibly used for the neural control of locomotion. To this end, a system with a double pendulum as a simplified model of the locomotor control and a model of bipedal locomotion were employed and analyzed in detail. In these models, a movement corresponding to the normal steady-state walking was realized as a stable limit cycle solution of the system. Unexpected external perturbations applied to the system can push the state point of the system away from its limit cycle, either outside or inside the basin of attraction of the limit cycle. Our mathematical analyses of the models suggested functional roles of the phase reset during walking as follows. Function 1: an appropriate amount of the phase reset for a given perturbation can contribute to relocating the system's state point outside the basin of attraction of the limit cycle back to the inside. Function 2: it can also be useful to reduce the convergence time (the time necessary for the state point to return to the limit cycle). In experimental studies during walking of animals and humans, the reset of walking rhythm induced by perturbations was investigated using the phase transition curve (PTC) or the phase resetting curve (PRC) representing phase-dependent responses of the walking. We showed, for the simple double-pendulum model, the existence of the optimal phase control and the corresponding PTC that could optimally realize the aforementioned functions in response to impulsive force perturbations. Moreover, possible forms of PRC that can avoid falling against the force perturbations were predicted by the biped model, and they were compared with the experimentally observed PRC during human walking. Finally, physiological implications of the results were discussed.     

A quantitative model of walking incorporating central and peripheral influences

Using the experimental results of Cruse and Saxler (1980a, b) and other authors (Graham, 1972; Pearson, 1972; Bässler, 1977, 1979) a quantitative model is developed in order to describe the behaviour of the systems controlling the leg movements of a walking insect. The whole model consists of six subsystems each of which controls the movement of an individual leg. The single subsystem (Fig. 1) consists of a central part which can assume two modes (protraction, retraction) the transition between which can be controlled by sensory influence. The central part produces the reference input for a feedback loop which controls the leg position. The reference input is however also determined by influences from other subsystems. Four different types of such connections are assumed to exist between the subsystems. Two of these produce alternating (t1, t3), two others in phase coupling (t2, t4) between the subsystems to be connected. These connections can transfer information originating from the central part as well as from the periphery of other subsystems. The model is capable of describing either quantitatively or qualitatively the experimental results of Cruse and Saxler (1980a, b) (see Figs. 3 and 4). In addition it is capable of describing the results of other authors, e.g. the temporal leg coordination of the free walking animal (Graham, 1972).Supported by DFG (Cr 58/1)     

Signaling oscillations: Molecular mechanisms and functional roles

Mounting evidence shows that oscillatory activity is widespread in cell signaling. Here, we review some of this recent evidence, focusing on both the molecular mechanisms that potentially underlie such dynamical behavior, and the potential advantages that signaling oscillations might have in cell function. The biological processes considered include cellular differentiation and tissue maintenance, intermittent responses in pluripotent stem cells, and collective cell migration during wound healing. With the aid of mathematical modeling, we review recent examples in which delayed negative feedback has been seen to act as a unifying principle that underpins this wide variety of phenomena.     

Energy efficient walking with central pattern generators: from passive dynamic walking to biologically inspired control

Like human walking, passive dynamic walking—i.e. walking down a slope with no actuation except gravity—is energy efficient by exploiting the natural dynamics. In the animal world, neural oscillators termed central pattern generators (CPGs) provide the basic rhythm for muscular activity in locomotion. We present a CPG model, which automatically tunes into the resonance frequency of the passive dynamics of a bipedal walker, i.e. the CPG model exhibits resonance tuning behavior. Each leg is coupled to its own CPG, controlling the hip moment of force. Resonance tuning above the endogenous frequency of the CPG—i.e. the CPG’s eigenfrequency—is achieved by feedback of both limb angles to their corresponding CPG, while integration of the limb angles provides resonance tuning at and below the endogenous frequency of the CPG. Feedback of the angular velocity of both limbs to their corresponding CPG compensates for the time delay in the loop coupling each limb to its CPG. The resonance tuning behavior of the CPG model allows the gait velocity to be controlled by a single parameter, while retaining the energy efficiency of passive dynamic walking.     

Asymmetries in the sense organs and central nervous system of the squid Histioteuthis

The visual system of Histioteuthis is markedly asymmetrical, in that the eyes and optic lobes are considerably larger on the left side, and the lens of the left eye is often yellower in colour than that of the right eye. At the histological level, the rhabdomes of the retinas of both eyes show the usual rectilinear pattern typical of cephalopods. Unlike other species described, however, the orientation of the pattern is not uniform over the retina. The optic lobes are well developed on both sides, again following the typical squid pattern, although the plexiform and inner granular layers are thicker on the left side. In life it is likely that the animals orient at an oblique angle with the arms downward, and the left eye pointing upwards and the right eye downwards, and the asymmetries of the visual system are probably related to this posture. No corresponding asymmetries in the statocysts or other parts of the central nervous system have, however, been detected     

The sense of consciousness

I propose that consciousness might be understood as the property of a system that functions as a sense in the biological meaning of that term. The theory assumes that, as a complex system, the sense of consciousness is not a fixed structure but implies structure with variations and that it evolved, as many new functions do, through the integration of simpler systems. The recognized exteroceptive and enteroceptive senses provide information about the organism's environment and about the organism itself that are important to adaptation. The sense of consciousness provides information about the brain and thus about the organism and its environment. It senses other senses and processes in the brain, selecting and relating components into a form that "makes sense"-where making sense is defined as being useful to the organism in its adaptation to the environment. The theory argues that this highly adaptive organizing function evolved with the growing complexity of the brain and that it might have helped resolve discrepancies created at earlier stages. Neural energies in the brain that are the input to the sense of consciousness, along with the processing subsystem of which they are a part, constitute the base of consciousness. Consciousness itself is an emergent effect of an organizing process achieved through the sense of consciousness. The sense of consciousness thus serves an organizing function although it is not the only means of organization in the brain. Its uniqueness lies in the character of the organization it creates with consciousness as a property of that organization. The paper relates the theory to several general conceptions-interactionism, epiphenomenalism and identity theory-and illustrates a number of testable hypotheses. Viewing consciousness as a property of a sense provides a degree of conceptual integration. Much of what we know about the evolution and role of the conventionally recognized senses should help us understand the evolution and role of the sense of consciousness, and of consciousness itself.     

Effect of nordic walking on functional ability and blood pressure in elderly women

The aim of the study was to analyze the effects produced by the use of experimental program (Nordic polewalking) on functional abilities in elderly women. Three-month polewalking led to reduction in the pulse rate at rest, diastolic and systolic blood pressure at the level of significance of 0.01 (p = 0.000). Polewalking improved the values of fitness index (FITIND) and maximal oxygen consumption (VO2max) at the level of significance of 0.01 (p = 0.000). On final measurement, three variables, i.e. pulse rate at rest - HRR (E = 73.42 vs. C = 79.68), systolic blood pressure - BPS (E = 118.42 vs. C = 123.65) and diastolic blood pressure - BPD (E = 79.04 vs. C = 83.54), showed lower results in experimental group compared with control group. On final measurement, experimental group showed higher values of the FITIND (E = 81.79 vs. C = 62.66) and VO2max (E = 21.83 vs. C = 16.81) variables as compared to control group. Accordingly, such a moderate physical activity, which is not too vigorous yet intensive enough to induce favorable changes, appears to be recommendable for elderly women. The present study included 60 women from the Novi Sad community, mean age 58.5 +/- 6.90 years, mean body mass 70.9 +/- 15.32 kg and mean body height 164.8 +/- 7.24 cm. Study population was divided into two groups of 30 subjects: experimental (E) group and control (C) group. The experimental program was performed three times a week for three months. The Nordic walking program was so designed for the performers to be in the aerobic work zone throughout the exercise. Nordic walking with poles was performed over three months. Study results revealed functional abilities of the study women to have modified during the longitudinal process.     

The role of the central nervous system functional condition in formation of responses of visual analyser's peripheral and central parts

Nembutal was shown to inhibit formation of early receptor potentials, "a"-wave and oscillatory potential of the retina (ERG), as well as the oscillatory potentials of the superior colliculus and visual cortex. The first oscillatory potential of the retina alone was enhanced. Aminazin exerted rather an opposite effect related, probably, to facilitation of the retina and central structures due to elimination of a constant inhibitory effect of the brain stem RF.     

The transmission sense of information

Biologists rely heavily on the language of information, coding, and transmission that is commonplace in the field of information theory developed by Claude Shannon, but there is open debate about whether such language is anything more than facile metaphor. Philosophers of biology have argued that when biologists talk about information in genes and in evolution, they are not talking about the sort of information that Shannon’s theory addresses. First, philosophers have suggested that Shannon’s theory is only useful for developing a shallow notion of correlation, the so-called “causal sense” of information. Second, they typically argue that in genetics and evolutionary biology, information language is used in a “semantic sense,” whereas semantics are deliberately omitted from Shannon’s theory. Neither critique is well-founded. Here we propose an alternative to the causal and semantic senses of information: a transmission sense of information, in which an object X conveys information if the function of X is to reduce, by virtue of its sequence properties, uncertainty on the part of an agent who observes X. The transmission sense not only captures much of what biologists intend when they talk about information in genes, but also brings Shannon’s theory back to the fore. By taking the viewpoint of a communications engineer and focusing on the decision problem of how information is to be packaged for transport, this approach resolves several problems that have plagued the information concept in biology, and highlights a number of important features of the way that information is encoded, stored, and transmitted as genetic sequence.