1 Zoologisches Institut, Universität zu
Köln, 50923 Cologne; 2 Fachbereich
Biologie, Universität Kaiserslautern, 67653 Kaiserslautern;
and 3 Chamissostrasse 16, 70193 Stuttgart, Germany
Akay, ...Turgay,
Ulrich Bässler,
Petra Gerharz, and
Ansgar Büschges.
The Role of Sensory Signals From the Insect Coxa-Trochanteral
Joint in Controlling Motor Activity of the Femur-Tibia Joint. J. Neurophysiol. 85: 594-604, 2001. Interjoint coordination in multi-jointed
limbs is essential for the generation of functional locomotor patterns.
Here we have focused on the role that sensory signals from the
coxa-trochanteral (CT) joint play in patterning motoneuronal activity
of the femur-tibia (FT) joint in the stick insect middle leg. This
question is of interest because when the locomotor system is active,
movement signals from the FT joint are known to contribute to
patterning of activity of the central rhythm-generating networks
governing the CT joint. We investigated the influence of femoral
levation and depression on the activity of tibial motoneurons. When the locomotor system was active, levation of the femur often induced a
decrease or inactivation of tibial extensor activity while flexor motoneurons were activated. Depression of the femur had no systematic influence on tibial motoneurons. Ablation experiments revealed that
this interjoint influence was not mediated by signals from movement
and/or position sensitive receptors at the CT joint, i.e., trochanteral
hairplate, rhombal hairplate, or internal levator receptor organ.
Instead the influence was initiated by sensory signals from a field of
campaniform sensillae, situated on the proximal femur (fCS). Selective
stimulation of these fCS produced barrages of inhibitory postsynaptic
potentials (IPSPs) in tibial extensor motoneurons and activated tibial
flexor motoneurons. During pharmacologically activated rhythmic
activity of the otherwise isolated mesothoracic ganglion (pilocarpine,
5 × 10 4 M),
deafferented except for the CT joint, levation of the femur as well had
an inhibitory influence on tibial extensor motoneurons. However, the
influence of femoral levation on the rhythm generated was rather labile
and only sometimes a reset of the rhythm was induced. In none of the
preparations could entrainment of rhythmicity by femoral movement be
achieved, suggesting that sensory signals from the CT joint only weakly
affect central rhythm-generating networks of the FT joint. Finally, we
analyzed the role of sensory signals from the fCS during walking by
recording motoneuronal activity in the single middle leg preparation
with fCS intact and after their removal. These experiments showed that
fCS activity plays an important role in generating tibial motoneuron
activity during the stance phase of walking.
Bath application of the muscarinic agonist pilocarpine onto the deafferented stick insect thoracic nerve cord induced long-lasting rhythmic activity in leg motoneurones. Rhythmicity was induced at ...concentrations as low as 1x10(-4) mol l-1 pilocarpine. The most stable rhythms were reliably elicited at concentrations from 2x10(-3) mol l-1 to 5x10(-3) mol l-1. Rhythmicity could be completely abolished by application of atropine. The rhythm in antagonistic motoneurone pools of the three proximal leg joints, the subcoxal, the coxo-trochanteral (CT) and the femoro-tibial (FT), was strictly alternating. In the subcoxal motoneurones, the rhythm was characterised by the retractor burst duration being correlated with cycle period, whereas the protractor burst duration was almost independent of it. The cycle periods of the rhythms in the subcoxal and CT motoneurone pools were in a similar range for a given preparation. In contrast, the rhythm exhibited by motoneurones supplying the FT joint often had about half the duration. The pilocarpine-induced rhythm was generated independently in each hemiganglion. There was no strict intersegmental coupling, although the protractor motoneurone pools of the three thoracic ganglia tended to be active in phase. There was no stereotyped cycle-to-cycle coupling in the activities of the motoneurone pools of the subcoxal joint, the CT joint and the FT joint in an isolated mesothoracic ganglion. However, three distinct 'spontaneous, recurrent patterns' (SRPs) of motoneuronal activity were reliably generated. Within each pattern, there was strong coupling of the activity of the motoneurone pools. The SRPs resembled the motor output during step-phase transitions in walking: for example, the most often generated SRP (SRP1) was exclusively exhibited coincident with a burst of the fast depressor trochanteris motoneurone. During this burst, there was a switch from subcoxal protractor to retractor activity after a constant latency. The activity of the FT joint extensor motoneurones was strongly decreased during SRP1. SRP1 thus qualitatively resembled the motoneuronal activity during the transition from swing to stance of the middle legs in forward walking. Hence, we refer to SRPs as 'fictive step-phase transitions'. In intact, restrained animals, application of pilocarpine also induced alternating activity in antagonistic motoneurone pools supplying the proximal leg joints. However, there were marked differences from the deafferented preparation. For example, SRP1 was not generated in the latter situation. However, if the ipsilateral main leg nerve was cut, SRP1s reliably occurred. Our results on the rhythmicity in leg motoneurone pools of deafferented preparations demonstrate central coupling in the activity of the leg motoneurones that might be incorporated into the generation of locomotion in vivo.
Structure function measurements and kinematic reconstruction at HERA Bassler, Ursula; Bernardi, Gregorio
Nuclear instruments & methods in physics research. Section A, Accelerators, spectrometers, detectors and associated equipment,
05/1999, Letnik:
426, Številka:
2
Journal Article
Recenzirano
Odprti dostop
The procedure used for structure function measurements at HERA is briefly described and related to the properties of kinematic reconstruction. The reconstruction methods of the inclusive deep ...inelastic scattering variables are reviewed and their sensitivity to the energy and angle miscalibrations are discussed in detail. New prescriptions are introduced and related to the standard methods in order to optimize the
F
2 structure function measurement over the widest kinematic range, both in the low
x, low
Q
2 and in the high
x, high
Q
2 regions. The prospects for the future high
Q
2 studies are briefly discussed.
Higgs boson studies at the Tevatron Agnew, J. P.; Annovi, A.; Bandurin, D. V. ...
Physical review. D, Particles, fields, gravitation, and cosmology,
09/2013, Letnik:
88, Številka:
5
Journal Article
Recenzirano
Odprti dostop
We combine searches by the CDF and DO Collaborations for the standard model Higgs boson with mass in the range 90- 200 GeV /c super(2) produced in the gluon-gluon fusion, WH, ZH, ttH, and vector ...boson fusion processes, and decaying in the H arrow right bb, H arrow right W super(+)W super(-), H arrow right ZZ, H arrow right tau super(+)tau super(-), and H arrow right gamma gamma modes. The data correspond to integrated luminosities of up to 10 fb super(-1) and were collected at the Fermilab Tevatron in pp collisions at radicals = 1.96 TeV. The searches are also interpreted in the context of fermiophobic and fourth generation models. We observe a significant excess of events in the mass range between 115 and 140 GeV /c super(2). The local significance corresponds to 3.0 standard deviations at mH = 125 GeV /c super(2), consistent with the mass of the Higgs boson observed at the LHC, and we expect a local significance of 1.9 standard deviations. We separately combine searches for H arrow right bb, H arrow right W super(+)W super(-), H arrow right tau super(+)t super(-), and H arrow right gamma gamma . The observed signal strengths in all channels are consistent with the presence of a standard model Higgs boson with a mass of 125 GeV /c super(2).
In a form of top-down analysis, the femur-tibia control system of stick insects is investigated. Open-loop experiments show that it is mainly velocity-sensitive with an extremely low ...velocity-threshold, that it possesses a very high gain and that it has only a very small phase reserve and thus works close to instability. The closed-loop system generates catalepsy. The system consists of a single sense organ with approximately 80 sense cells with known characteristics, a small number of interneurones (mainly non-spiking ones) and a small number of motor neurones. The characteristics of the whole system can quantitatively be attributed to the characteristics of its elements. The gain of the loop is state-dependent and the system is 'switched off' during active movements and replaced by a control system with different attributes. It is discussed that most of the characteristics of this system are, at least qualitatively, similar to joint control systems in other animals. Because the described system can be more easily investigated than other systems (especially in vertebrates) it can serve as a model against which more complicated joint control loops may be compared.
The extremely slow return movements observed in stick insects (phasmids) after imposed changes in posture are termed catalepsy. In the literature, catalepsy is treated as a behavioural component of ...the twig mimesis observed in walking stick insects. It is produced by the high gain of the velocity-sensitive component of the relevant joint control systems and by the non-linear dependency of its time constant on movement velocity. The high gain, in turn, causes the system to work close to instability, and this may have driven the evolution of gain control mechanisms. Although these statements represent plausible assumptions, based on correlated occurrence, they remain largely hypothetical like many ideas concerning evolutionary tendencies. To test these hypotheses, we studied catalepsy and the relevant properties of the femur-tibia control system in the middle and hind legs of Prosarthria teretrirostris.cf. Prosarthria teretrirostris is a proscopiid closely related to grasshoppers and locusts. With its slender, green-to-brown body and legs, it shows clear morphological twig mimesis, which has evolved independently of the well-known twig mimesis in stick insects. The animals show clear catalepsy. The main properties of femur-tibia joint control are remarkably similar between proscopiids and stick insects (e.g. the marked sensitivity to movement velocity rather than to joint position and the non-linear dependency of the time constants of response decay on movement velocity), but there are also important differences (habituation and activity-related mechanisms of gain control are absent). Together, these results validate the main concepts that have been developed concerning the neural basis and evolution of catalepsy in stick insects and its relationship to twig mimesis, while demonstrating that ideas on the role of habituation and gain control should be refined.
It is shown that the low-pass filter characteristics of the musclejoint system of the femurtibia joint of the stick insect Cuniculina impigra result from co-contraction of the extensor and flexor ...tibiae muscles. The most distal region of the extensor muscle, which contains a high percentage of slow muscle fibres, is involved in this co-contraction. This conclusion results from the following evidence. (1) Inertial and friction forces do not affect the characteristics of the low-pass filter of the musclejoint system. (2) There is some co-contraction of the extensor and flexor muscles during sinusoidal stimulation of the femoral chordotonal organ at high stimulus frequencies. Both muscles generate tonic forces that increase with increasing stimulus frequency and also increase with time from the beginning of stimulation until a plateau is reached. (3) For the extensor muscle, this tonic force is produced by its most distal portion only. (4) Electrical stimulation of the common inhibitory motoneurone (CI1) reduces the tonic force generated in this most distal portion of the extensor muscle. Therefore, CI1 stimulation reduces the amplitude of tibial movement in response to sinusoidal stimulation of the femoral chordotonal organ at stimulus frequencies below 0.5 Hz (over this frequency range, the tibial movement amplitude is a function of the force amplitude produced by the whole extensor muscle and there is no co-contraction), but at chordotonal organ stimulus frequencies of 1 Hz and above, CI1 stimulation increases the tibial movement amplitude (in this case, movement amplitude is limited by the degree of co-contraction of the extensor and flexor muscles). With repeated chordotonal organ stimulation at higher stimulus frequencies, the tibial movement amplitude steadily decreases. This must be a consequence of increasing levels of co-contraction of the extensor and flexor muscles, since at low stimulus frequencies (no co-contraction) there is no reduction in movement amplitude during repeated stimulations. It is concluded that co-contraction of the extensor and flexor tibiae muscles prevents instability in the reflex loop in spite of the high gain necessary for the generation of catalepsy. Therefore, the mechanism described can be considered to be an adaptation to the ecological niche occupied by this animal. The contribution of the distal part of the extensor muscle to this system can be switched off by the CI1 during active movements.
In orthopteran insects, neural networks for joint control exhibit different characteristics due to behavioural specializations. We investigated whether these differences are generated purely by the ...neuronal networks, or whether characteristics of the muscles or joint architecture (musclejoint system) are also involved in these behavioural specializations. We compared the properties of the muscle system moving the femurtibia joint of the middle and hindleg of three species, Carausius morosus, Cuniculina impigra and Locusta migratoria. Four aspects were analysed for the tibial extensor muscle: (i) the frequency-dependence of motoneuronal activity in response to sinusoidal stimulation of the femoral chordotonal organ (fCO), (ii) the muscle structure, (iii) the innervation pattern of the muscle and (iv) the histochemical properties of the muscle fibres. These aspects were compared with the filter characteristics of the open-loop femurtibia control system and of the musclejoint system involved. Whereas in both phasmid species (Carausius morosus and Cuniculina impigra) the motoneuronal activity steadily increases with sinusoidal stimulation of the fCO in the frequency range 0.015 Hz, in Locusta migratoria there is a decrease in motoneuronal activity between 0.01 and 0.3 Hz. The muscle structure is basically similar in all three species, as the number of singly innervated muscle fibres (supplied by the fast extensor tibiae motor neurone, FETi) decreases from proximal to distal. The number of triply innervated fibres supplied by the FETi, the slow extensor tibiae (SETi) and the common inhibitor 1 (CI1) is maximal in the middle of the muscle, and the number of dually innervated fibres (supplied by SETi, CI1) increases from proximal to distal. Differences between the locust and the two phasmid species exist in the distal portion of the muscle. The phasmid extensor tibiae muscle contains a morphologically distinct bundle of muscle fibres, not present in the locust, which is mostly dually innervated and which is larger in Cuniculina impigra. Similar results were obtained for the histochemical characterisation of the muscle fibres as revealed from their staining for myofibrillar ATPase activity. The number of histochemically identified fast fibres decreased from proximal to distal, while the number of slow fibres increased. In Carausius morosus and Locusta migratoria, the percentage of slow fibres increased by up to 6070 % at the distal end, while this increase was to almost 100 % in Cuniculina impigra. Apparently, the larger this distal region and the higher the percentage of slow, dually innervated fibres in it, the lower is the upper corner frequency (the stimulus frequency at which the joint control system produces a movement with 70 % of its maximal response amplitude) of the musclejoint system. In summary, it appears that the upper corner frequency of the open-loop system in Locusta migratoria (<0.05 Hz) results at least in part from properties of the neuronal joint control network, but in Carausius morosus (0.51.0 Hz) and Cuniculina impigra (0.10.2 Hz) it results from the upper corner frequency of the musclejoint system.
We present the results of the combination of searches for the standard model Higgs boson produced in association with a W or Z boson and decaying into bb using the data sample collected with the D0 ...detector in pp collisions at √s = 1.96 TeV at the Fermilab Tevatron Collider. We derive 95% C.L. upper limits on the Higgs boson cross section relative to the standard model prediction in the mass range 100 GeV ≤ M(H) ≤ 150 GeV, and we exclude Higgs bosons with masses smaller than 102 GeV at the 95% C.L. In the mass range 120 GeV ≤ M(H) ≤145 GeV, the data exhibit an excess above the background prediction with a global significance of 1.5 standard deviations, consistent with the expectation in the presence of a standard model Higgs boson.
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