Identification and characterization of gibberellin-insensitive mutants selected from among dwarf mutants of rice

In rice, many dwarf mutants have been isolated and characterized. We have investigated the relationship between dwarfism and the gibberellin (GA)-mediated control of physiological processes. Twenty-three rice cultivars and mutants (9 normal, 3 semi-dwarf, 11 dwarf) were analyzed in terms of two GA-mediated processes, namely, elongation of shoots and production of -amylase activity in the endosperm. As a result, we identified four different groups (groups N, T, D and E). Two-dimensional plotting of the extent of induction of -amylase in the endosperm versus the extent of enhancement of shoot elongation upon treatment with exogenous gibberellic acid (GA₃) provided a useful method for the rapid allocation of large numbers of dwarf mutants of rice to the various groups. Members of group N (normal type), which included all normal cultivars and semi-dwarf mutants, showed a slight increase in elongation of shoots and a remarkable increase in production of -amylase with the application of GA₃ during germination. All of the dwarf mutants were classified as being members of the other three groups. Members of group T (Tan-ginbozu type), including three dwarf mutants, were highly responsive to exogenous GA₃ in terms of elongation of shoots and production of -amylase, with associated lower levels of endogenous GA. In contrast, members of the other three groups, including group N, had normal levels of endogenous GAs. Members of group D (Daikoku type) were only slightly responsive to exogenous GA₃, an indication that they are GA-insensitive mutants. Members of group E (Ebisu type) had responses to GA₃ similar to those of group N, not only in terms of elongation of shoots but also in terms of -amylase production, an indication that they are dwarf mutants that can be considered as neither GA-deficient nor GA-insensitive mutants. We also examined a GA-insensitive mutant selected from among 19 near-isogenic dwarf lines of Shiokari, and we concluded that the d-1 gene is associated with the phenotype of GA-insensitive dwarf mutants.     

Walking on a ‘peg leg’: extensor muscle activities and sensory feedback after distal leg denervation in cockroaches

Previous studies in insects demonstrated that leg coordination changes following complete ablation of distal limb segments. However, normal coordination was restored when small peg leg prostheses were attached to leg stumps to permit substrate contact. We have adapted this paradigm to preserve appropriate leg mass and inertia by severing all nerves and muscle tendons in the femur of the cockroach hind leg and converting the animals own limb into a peg leg. Recordings of muscle activities and leg movements before and after denervation showed that: (1) the peg leg is actively used in walking and regular bursts occur in motoneurons to leg extensor muscles; (2) driving of motoneuron activity is sufficient to produce fictive bursting in a muscle whose tendon (apodeme) is cut in the ablation; and (3) similar motoneuron activities are found in walking on an oiled glass surface, when the effects of body weight and mechanical coupling are minimized. When distal segments were completely severed in these preparations, leg use and muscle bursting were disrupted but could be restored if the stumps were pressed against the substrate. These results support the hypothesis that feedback from receptors in proximal leg segments indicating forces allows for active leg use in walking.     

Vom femoralen Chordotonalorgan gesteuerte Reaktionen bei der Stabheuschrecke Carausius morosus: Messung der von der Tibia erzeugten Kraft im aktiven und inaktiven Tier

In the Introduction (A) there is a list of unsolved problems concerning the role of the femoral chordotonal organ. A method to solve these problems by measuring the force at the distal end of the tibia during stimulation of the femoral chordotonal organ is described in (B). The step-response in inactive animals (C) is similar to that of the free-moving tibia. After an active movement caused by touching the abdomen the amplitude of the flexion-force is always higher than before. In (D) a method is described to measure the amplification of the control-system in intact animals. With this method it is verified, that the flexion-force produced by a distinct stimulus is higher after active movements caused by touching the abdomen. But this force is lower after spontaneous active movements caused by darkening the room (Fig. 2). Therefore one must assume, that there are two different types of activity: spontaneous activity and activity after a disturbance. In the frequency-response of the inactive animal (F) (Figs. 4 and 5) the amplitude of the force decreases with increasing frequency at a constant amplitude of stimulus. The phase-shift between reaction and stimulus is much smaller than with the free-moving tibia. Therefore, the large phase-shift as well as the strong decrease of the reaction-amplitude near 1 Hz observed in free-moving tibias (1972b) is mainly due to the mechanical attributes of the system. In Section (F) the receptor-apodeme is sinusoidally moved during active movements of intact and decerebrated animals. As with the free-moving tibia no reaction can be observed during active movements at that phase position for which the response occurs in inactive animals. Instead of this inactive response there is another response, called active with a phase-shift of about 180°. At the end of an active period the active and the inactive response can be observed simultaneously (Figs. 7 and 10). The amplitude of the active response decreases, and the amplitude of the inactive response increases from cycle to cycle. In decerebrated animals there are normally several minutes from the exclusively active response to the exclusively inactive response without a further increase in amplitude. In intact animals this transition takes only a few seconds. Step-stimuli during active movements (G) show, that in active animals stretching the chordotonal organ causes a flexion of the femor-tibia-joint. Releasing the chordotonal organ does not produce any reaction. Moving the receptor-apodeme in active animals influences the contralateral leg significantly only in middle legs (H). These legs tend to move within the same phase position as the stimulated leg. Moving the receptor-apodeme in a middle leg has no influence on the ipsilateral hind leg, but a weak influence on the ipsilateral front leg, which tends to move within the same phase position as the middle leg. In the discussion (I) a hypothesis is presented according to which the active response is a mechanism for adapting the leg movement to a surface which suddenly gives way (I 5). The influence on the contralateral middle leg seems to be a part of this mechanism (I 6). This reaction has nothing to do with the coordination of leg movements in walk (I 7). The feed-back systems which control the distance between the body and the walking surface may be inactive during walking (I 8), but those systems which control the forward movement of the body must be active. Since the feed-back system of the Kniesehnen-reflex controls predominantly the body-ground-distance it seems likely that it is normally inactive during walking.     

Two major groups of colicin factors: Their evolutionary significance

Summary Eleven colicin factors have been placed in two groups defined by a number of physiological criteria such as the effect of the host recA (recombination-proficiency) allele on colicin titres and the maximum number of copies of the colicin factor per chromosome. The fundamental difference between the two groups may lie in the molecular weight of the plasmid DNA: one group is about 5x10⁶, the other about 70x10⁶. The colicins specified by members of each group are also related. Colicin factors within the same group may therefore resemble each other because they are descended from the same ancestral plasmid which was either EK-like of low molecular weight or BIV-like of high molecular weight.Aided by grants from the Science Research Council and the Central Research Fund of the University of London.     

Influence of loading parallel to the body axis on the walking coordination of an insect

It is often reported in the early literature that insects walk with the legs protacting in diagonal pairs rather than the triplet of three legs associated with the tripod step pattern. The diagonal pattern implies that legs of the same segment have a phase relationship significantly different from 0.5. Such a pattern of leg recovery has been demonstrated quantitatively for the stick insect (Graham, 1972). Such patterns occur in several insects and systematic asymmetry can even be detected in the earliest quantitative study on cockroaches (Hughes, 1957) when the animals are walking slowly. More recently Spirito and Mushrush (1979) have reported systematic deviations from a phase of 0.5 similar to those observed in stick insects. Asymmetry has also been quantitatively demonstrated in Katydids (Graham, 1978) and has recently been observed in Mantid walking (Thomson, personal communication). This phenomenon seems to be a general characteristic of slow walking coordination in insects. In stick insects asymmetry only becomes obvious in gait II at slow speeds although there can be systematic differences in ipsilateral coordination on right and left sides even at the highest speeds in this gait (Graham, 1972).     

Measurements of electrical potential differences across yeast plasma membranes with microelectrodes are consistent with values from steady-state distribution of tetraphenylphosphonium inPichia humboldtii

Summary Electrical potential differences across the plasma membrane () of the yeastPichia humboldtii were measured with microelectrodes (filled with 0.1m KCl) inserted into cells immobilized in microfunnels. The registered signals were reproducible and stable for several minutes. On attainment of stable reading for the specific membrane resistanceR _sp was determined by applying square-current pulses to the preparation. Both andR _sp were pH dependent and displayed equal but opposite deflection, reaching its maximal value of –88±9 mV (n=13) andR _sp its minimal value of 10 k·cm² (maximal conductance) at pH 6.5. Uncouplers and the polyene antibiotic nystatin depolarized the cells, decreasing to –21±15 mV (n=10) with concomitant decrease ofR _sp. Comparison of values from microelectrode measurements with those calculated from the steady-state distribution of tetraphenylphosphonium ions agreed within 10 mV under all physiological conditions tested, except at pH values above 7.0. During microelectrode insertion transient voltage signals (a few msec long) were detected by means of an oscilloscope. These voltage signals were superimposed on the stable recordings described above. These short voltage signals disappeared in uncoupled cells. The closely related values obtained by two independent methods (direct measurements with microelectrodes and calculation from steady-state distribution of a lipophilic cation) provide evidence that these values reffect the true membrane potential of intact cells.     

Role of the Na,K-ATPaseβ-subunit in the cellular accumulation and maturation of the enzyme as assessed by glycosylation inhibitors

Summary No functional role could yet be established for the glycosylated -subunit of the Na,K-ATPase. In this study, we describe the intracellular processing of the -subunit as a glycoprotein in toad bladder cells and the consequences of its structural perturbation with glycosylation inhibitors on the cellular expression of the - and -subunits and on the structural and functional maturation of the enzyme. Controlled trypsinolysis of homogenates from pulse-labeled cells reveals that the -subunit is subjected to glycosylation-dependent structural rearrangements during its intracellular routing. Inhibition of correct terminal glycosylation of the -subunit with deoxynojirimycin or swainsonine has no effect on the trypsin sensitivity of the -subunit, its ability to perform cation-dependent conformation changes or the cellular Na,K-ATPase activity. Acquisition of core-sugars is sufficient for the enzyme to assume its catalytic functions. On the other hand, complete inhibition of glycosylation with tunicamycin leads to a destabilization of both the - and the -subunits as judged by their higher trypsin sensitivity. In addition, tunicamycin treatment results in a decrease of the amount of newly synthesized - and -subunit indicating that a glycoprotein, possibly the -subunit itself, plays a role in the efficient accumulation of the -subunit in the endoplasmic reticulum.     

A self-organizing model of walking patterns of insects

It is well known that the motor systems of animals are controlled by a hierarchy consisting of a brain, central pattern generator, and effector organs. An animal's walking patterns change depending on its walking velocities, even when it has been decerebrated, which indicates that the walking patterns may, in fact, be generated in the subregions of the neural systems of the central pattern generator and the effector organs. In order to explain the self-organization of the walking pattern in response to changing circumstances, our model incorporates the following ideas: (1) the brain sends only a few commands to the central pattern generator (CPG) which act as constraints to self-organize the walking patterns in the CPG; (2) the neural network of the CPG is composed of oscillating elements such as the KYS oscillator, which has been shown to simulate effectively the diversity of the neural activities; and (3) we have introduced a rule to coordinate leg movement, in which the excitatory and inhibitory interactions among the neurons act to optimize the efficiency of the energy transduction of the effector organs. We describe this mechanism as the least dissatisfaction for the greatest number of elements, which is a self-organization rule in the generation of walking patterns. By this rule, each leg tends to share the load as efficiently as possible under any circumstances. Using this self-organizing model, we discuss the control mechanism of walking patterns.     

Visual control of turning responses to tactile stimuli in the crayfishProcambarus clarkii

Summary Specimens of the crayfishProcambarus clarkii turn to face in the direction of a brief tactile stimulus delivered to a walking leg. The control system that guides this directed behavior was investigated under closed-loop and open-loop conditions. The accuracy of turns exhibited in these experiments was compared to baseline accuracy established by animals restrained from forward and backward walking but allowed to rotate in the yaw plane. Procambarus clarkii individuals deprived of visual feedback tended to undershoot the target angle. Response accuracy increased when a uniform field of stripes moved across the visual field in accordance with the turning movements of the animal. Response accuracy did not match the accuracy observed under baseline conditions, however, unless the responding animal encountered a novel visual image, such as the silhouette of a crayfish, in the moving visual field.Visual feedback thus influences the accuracy of turning in crayfish in two important ways. Movement of stripes across the visual field of a crayfish feeds back positively and promotes rapid turning during the initial phase of a response. This effect obtains regardless of the direction or rate of movement of the stripes in the visual field. The appearance of a novel image in the visual field feeds back negatively to inhibit at least partially further turning. Feedback from the visual system appears to fine tune basic turning movements initiated by a tactile stimulus and crudely directed according to that input. Turning behavior in the crayfish resembles in this respect compensatory eye movements in the lobster and escape responses in a number of arthropods.Neural mechanisms that may explain the experimental results are discussed with particular emphasis on the possibility of interaction between voluntary turning responses and optomotor reactions.     

Analysis of myelin proteolipid protein and F0ATPase subunit 9 in normal andjimpy CNS

Membrane fractions and chloroform-methanol (C-M) extracts ofjimpy (jp) and normal CNS at 17–20 days were examined by immunoblot and sequence analysis to determine whether myelin proteolipid protein (PLP) or DM-20 could be detected in jp CNS. No reactivity was detected in jp samples with several PLP antibodies (Abs) except with one Ab to amino acids 109–128 of normal PLP. Proteins in the immunoreactive bands 26 Mr comigrating with PLP were sequenced for the first 10–12 residues. A sequence corresponding to PLP was found in normal CNS, as expected, but not in the band from jp CNS. Our results provide no evidence for an aberrant form of PLP in jp CNS at 17–20 days. This and other studies suggest that the abnormalities in jp brain are not due to toxicity of the mutant jp PLP/DM-20 proteins. Interestingly, a sequence identical to the amino terminus of the mature proton channel subunit 9 of mitochondrial F₀ ATPase was detected in the immunoreactive bands 26 Mr in both normal and jp samples. This identification was supported by reactivity with an Ab to the F₀ subunit and by labeling with dicyclohexylcarbodiimide (DCCD). In contrast to PLP isolated from whole CNS, PLP isolated from myelin was devoid of F₀ subunit 9 based on sequence analysis and lack of reactivity with an Ab to the F₀ subunit, yet still reacted with DCCD. This finding rules out the possibility that contaminating F₀ ATPase gives rise to the DCCD binding exhibited by PLP and confirms the possibility that PLP has proton channel activity, as suggested by Lin and Lees (1,2).Abbreviations used Ab antibody - CM conditioned medium - C M chloroform-methanol - DCCD dicyclohexylcarbodiimide - jp jimpy - Mr mobility (apparent m.w×10^–3) - PLP proteolipid protein - PVDF polyvinylidene difluoride     

The hypothalamo-hypophysial system in acipenseridae

Summary The neurohypophysis of sexually mature male and female Acipenser güldenstädti Brandt and Acipenser stellatus Pallas was studied light and electron microscopically. The recessus hypophysei lined with ependymal cells of two main types, narrow and wide, are in the center of the neurohypophysial roots. Processes from both cell types run radially to the basement membrane of the connective tissue layers abutting on the hypophysial intermediate lobe. Protrusions penetrating deep into the recessus hypophyseus are found in the apical parts of the wide cells. Pituicytes are rare in the neurohypophysis. The ultrastructure of both ependymal cell types and of the pituicytes is described. Nonmyelinated Gomori-positive (peptidergic) neurosecretory A₁ and A₂ type fibres and their terminals containing elementary neurosecretory granules (1400–1800 Å and 1000–1400 Å respectively) are the main structural elements of the neurohypophysis. Some dark and single myelinated neurosecretory fibres have been found. The adrenergic fibres (type B) were described earlier (Polenov et al., 1972a). The structural peculiarities of the neurohypophysis are discussed in functional and comparative-morphological terms.     

Synthesis of two-dimensional human walking: a test of the <Emphasis Type="Italic">λ</Emphasis>-model

To test the -model version of the equilibrium point hypothesis both for feasibility and validity with respect to the control of terrestrial locomotion, we developed a two-dimensional, eleven-segment musculoskeletal model of the human body including 14 muscle-tendon complexes per leg, three-segment feet, and a physiologically based model of foot-ground interaction. Human walking was synthesized by numerical integration of the coupled muscle-tendon and rigid body dynamics. To this end a control algorithm based on the -model was implemented in the model providing muscle stimulation patterns that guaranteed dynamically stable walking including a balanced trunk. Thus, the timing of the movement is not preset by a central pattern generator but emerges from the interaction of the musculoskeletal system with the control algorithm. The control parameters were found in a trial-and-error approach. The feedforward part of the control scheme consists of just two target configurations each of which is composed of a set of one nominal length per muscle (-model). Variation of gravity reveals that (1) the synthesized walking patterns are close to ballistic walking and (2) this muscularly induced natural walking can only be initiated and maintained in the range between about a tenth and three times earth-bound gravity. Our walking patterns are robust both against parameter variations and shuffling of the swing leg. We discuss our model with respect to gravity scaling, speed control, feedback delay, and the terms equilibrium point hypothesis and central pattern generator.     

Somatic embryogenesis from cultured leaf segments of Zea mays

Basal leaf segments of 3 to 4 week old maize (Zea mays L.) seedlings plated on SH medium with 30 M dicamba produced embryogenic callus and/or somatic embryos. Histological evidence showed that some of the embryos arose directly from the explant. When leaf segments with embryos were transferred to MS medium with 1.0 M NAA, 1.0 M IAA, 2.0 M 2iP, and 60 g/l sucrose, the embryos germinated and the resulting seedlings could be established in culture tubes. These responses were obtained from three inbred lines, CHI31, S615, and S7.Abbreviations SH Schenk and Hildebrandt (1972) medium - MS Murashige and Skoog (1962) medium - dicamba 3,6-dichloro-o-anisic acid - IAA indole-3-acetic acid - NAA -naphthaleneacetic acid - 2iP 2-isopentyladenine     

Sensory innervation of the tentacles of the polychaete,Sabella pavonina

Summary Following observation of conical groups of stiff, but motile cilia on the tentacles of the branchial crown of Sabella pavonina, these were examined with the electron microscope. The bundles consist of about 40 unenclosed standard cilia supported by one or two primary sense cells with centrally directed axons of 0.1–0.2 diameter. Axons in the distal portions of the branchial crown occur in small bundles surrounded by a basement membrane. More centrally, glial elements appear and the nerves are surrounded by a collagenous sheath. The branchial nerve trunk shows similarities in organisation to other previously investigated annelid central nervous tissue in that the whole nerve is surrounded by a fibrous sheath central to which there is a layer of glial cells with processes penetrating a central neuropile. The 0.1–0.2 axons commonly occur in glial-enveloped groups of < 40 whilst other axons of larger and mixed diameter are found together.Each tentacle has two branchial nerves on the oral side, and each nerve gives rise to two small 75-axon branches running to each pinnule. The branchial nerves fuse to form the branchial nerve trunk running to the supra-oesophageal ganglia.Sections of the branchial nerves of the branchial crown at progressively more central levels show that the branchial nerve trunk contains enough axons of 0.1–0.2 diameter to account for all the sensory cells on the tentacles. This is taken as evidence for the sensory cells having axons terminating within the central nervous system and that there is no peripheral confluence or fusion of these afferent axons.     

Regeneration of Elaeagnus angustifolia from leaf segments of in vitro-derived shoots

Shoot regeneration was achieved from in vitro-produced leaves of Elaeagnus angustifolia L. Half-leaf explants from the terminal part of the shoot produced more shoots than explants from the basal part of the in vitro-derived shoots on agar-solidified WPM medium supplemented with 1 M benzyladenine (BA). In liquid medium of the same formulation, compact shoots that did not elongate were formed on the explants. Leaf cross-section explants (1 mm thick) produced shoots both on solid and liquid medium with 1 M BA, whereas again compact shoots were formed with 10 M BA. Further shoot development on these explants was promoted by their transfer to fresh solid medium containing 1 M BA and 1 M gibberellic acid (GA₃).Abbreviations BA benzyladenine - GA₃ gibberellic acid - WPM woody plant medium     

Disulfiram lowers Ca2+, Mg2+-ATPase activity of rat brain synaptosomes

The chronic administration of disulfiram (DS) to rats resulted in significant decrease of synaptosomal Ca²⁺, Mg²⁺-ATPase activity. In vitro studies indicated that DS (ID₅₀=20 M) produced a dose-dependent inhibition of Ca²⁺, Mg²⁺-ATPase. However, diethyldithio-carbamate, a metabolite of DS, failed to modify Ca²⁺, Mg²⁺-ATPase activity, implying that the decrease in ATPase activity in DS administered rats was due to the effect of parent compound. The DS-mediated inhibition (48%) of ATPase activity was comparable with a similar degree of inhibition (49%) achieved by treating the synaptosomal membranes with N-ethylmaleimide (ID₅₀=20 M) in vitro. Furthermore, the inhibition by DS was neither altered by washing the membranes with EGTA nor reversed by treatment with sulfhydryl reagents such as GSH or dithiothreitol. About 74% and 68% decrease of synaptosomal Ca²⁺, Mg²⁺-ATPase specific activity was observed when treated with DS (30 M) and EGTA (100 M) respectively. The remaining 25–30% of total activity is suggested to be of Mg²⁺-dependent ATPase activity. This indicates that both these drugs may act on a common target, calmodulin component that represents 70–75% of total Ca²⁺, Mg²⁺-ATPase activity. Therefore, DS-mediated modulation of synaptosomal Ca²⁺, Mg²⁺-ATPase activity could affect its function of maintaining intracellular Ca²⁺ concentration. This could contribute to the deleterious effects on CNS.     

Permeability of submandibular salivary glands in dogs to blood-borne horseradish peroxidase (HRP)

Summary Horseradish peroxidase (HRP) was administered to the submandibular glands of dogs by close-arterial bolus-type injections, and its localisation was examined histochemically by light and electron microscopy. The HRP became widespread in the interstices of the glands and reached many central acinar lumina via scattered localised parts of their tight junctional complexes. Reaction product was less often found in the lumina of demilunes, which suggested that the intercellular junctions there were less leaky. HRP was often found in sizeable spaces between myoepithelial cells and the underlying parenchymal cells; such large spaces have not been observed in this situation in other species. The possibility that permeability pathways may arise intermittently at different sites in the adhering mechanisms between the acinar cells is discussed.It is concluded that potential paracellular permeability pathways for macromolecules exist in these glands and, if the concentration gradient is sufficiently high, molecules even as large as those of HRP can to some extent permeate passively from the interstices to the saliva. In resting glands the principal permeability site is between the central acinar cells.Supported by Grants from the M.R.C. and the V.R.T. King's College HospitalWe wish to acknowledge the technical help of Mr. K.J. Davies and Mr. P.S.A. Rowley     

Staphylococcus aureus alpha-toxin-induced pores: Channel-like behavior in lipid bilayers and patch clamped cells

The conductance of pores induced by Staphylococcus aureus -toxin in Lettre cells has been compared to that in bilayers composed of synthetic lipids or Lettre cell membrane constituents. Previously described characteristics of toxin-induced conductance changes in lipid bilayers, namely rectification, voltage-dependent closure, and closure at low pH or in the presence of divalent cations (Menestrina, 1986) are displayed also in bilayers prepared from Lettre cell membranes and in patch clamped Lettre cells. It is concluded that endogenous proteins do not affect the properties of -toxininduced channels significantly and that the relative lack of ion channels in Lettre cells makes them ideal for studies of pore-forming toxins by the patch clamp technique.Dr. Sviderskaya is on leave of absence from the Physiology Institute, University of St. Petersburg, RussiaWe are grateful to Dr. J.P. Arbuthnott and Dr. K. Hungerer for gifts of S. aureus -toxin, to Dr. T.B. Bolton for collaboration with patch clamped cells and to Dr. J.M. Graham for help with the preparation of Lettre cell plasma membranes. This study was supported by the Cell Surface Research Fund, Medical Research Council, Science and Engineering Research Council, UNESCO (Molecular and Cell Biology Network) and The Wellcome Trust.     

Sensory control of leg movement in the stick insect Carausius morosus

Hind legs with crossed receptor-apodemes of the femoral chordotonal organ when making a step during walking often do not release the ground after reaching the extreme posterior position. After putting a clamp on the trochanter (stimulation of the campaniform sensilla) the leg is no longer protracted during walking. However, during searching-movements the same leg is moved very far forwards. The anatomical situation of the campaniform sensilla on the trochanter and the sensory innervation of the trochanter is described. After removal of the hair-rows and continuously stimulating the hair-plate at the thorax-coxa-joint the extreme anterior and posterior positions of the leg in walking are displaced in the posterior direction. Front and middle legs operated in this way sometimes do not release the ground at the end of retraction. In searching-movements the same leg is moved in a normal way. If only one side of a decerebrated animal goes over a step, then on the other side a compensatory effect is observed. The main source of this compensatory information appears to be the BF1-hair-plates. If the animal has to drag a weight the extreme anterior and posterior positions of the middle and hind legs are displaced in the anterior direction. Crossing the receptor-apodeme of the femoral chordotonal organ, when it causes the leg to remain in the protraction phase, displaces the extreme posterior position of the ipsilateral leg in front of the operated one in the posterior direction. Influences of different sources on the extreme posterior position can superimpose. A model is presented which combines both a central programme and peripheral sensory influence. The word programme used here means that it does not only determine the motor output but also determines the reactions to particular afferences. The fact that the reaction to a stimulus depends on the internal state of the CNS is also represented by the model.Supported by Deutsche Forschungsgemeinschaft