Sensory feedback in the control of mouthpart movements in the desert locust Schistocerca gregaria

ABSTRACT. When imposed movements were applied to one or both mandibles of the desert locust, Schistocerca gregaria , the other mouthparts moved in synchrony with the mandibles. This occurred in the presence or absence of food, and when the mandibles were driven at a higher or lower frequency than that seen during normal feeding. Electromyogram recordings from the mandibular closer muscles revealed bursts of activity at the same frequency as the imposed movement. This activity occurred during mandibular closing. Burst length was a function of driving wavelength. At low driving frequencies (less than 0.5 Hz), smaller bursts were seen prior to the longer closing burst; a series of similar small bursts was seen when the mandibles were held in the open position. When one mandible was driven, closer muscle activity was largely confined to that side. In the presence of food, however, activity was seen in both closer muscles. A possible mechanism for this is described. After destruction of the campaniform sensilla on the ventral surface of the mandibles, the bursts of activity in the mandibular closers, seen when the mandibles were held open, were replaced by continuous activity. This suggests that the function of these sensilla is to inhibit motor output to the closer muscles when the tension becomes high. When feeding on relatively incompressible food the closer muscle burst length increased, although chewing frequency did not alter. This effect was also produced by loading the mandibles artificially. A model for the feedback control of this behaviour is proposed.     

Preweaning feeding mechanisms in the rabbit.

Muscle contraction patterns and mandibular movements of infant rabbits during suckling and chewing were compared. Oral muscle activity was recorded by fine-wire electromyography, while jaw movements and milk bottle pressure were registered. Suckling and mastication have a comparable cycle duration and share a common pattern of oral muscle activity which consists of a succession of a jaw closer burst, during which the jaw closes and undergoes a power stroke (in mastication), a suprahyoid burst with a stationary or slightly opening jaw and a digastric burst with fast jaw opening (the power stroke of suckling). Compared to suckling, mastication shows decreased jaw opener activity, increased jaw closer activity, development of jaw closing activity in the lateral pterygoid, and increased asymmetry in the masseter by development of a new differentiated motor pattern on the working side. The study shows that the suckling motor pattern enables the infant rabbits to change to chewing with just a few modifications.     

Fictive chewing activity in motor neurons and interneurons of the suboesophageal ganglion of Manduca sexta larvae

Alternating antiphasic rhythmic activity was observed in opener and closer mandibular motor neurons in the isolated suboesophageal ganglion of the larva of Manduca sexta (Lepidoptera: Sphingidae). This was interpreted provisionally as fictive chewing; the pattern is similar to that seen in semiintact animals but of lower frequency. Additionally, a variety of associated rhythmic activities were observed in suboesophageal interneurons. These could be classified into several different physiological types by their activity patterns in relation to the chewing cycle. Some of these neurons can modulate the rhythm when injected with current. It seems likely that they are part of or associated with a central pattern generator circuit for chewing.Abbreviations A anterior - CEC circumoesophageal connective - Cl-MN closer motor neuron - IN interneuron - MdN mandibular nerve - MN motor neuron - O-MN opener motor neuron     

Muscle activity reduces soft-tissue resonance at heel-strike during walking

Muscle activity has previously been suggested to minimize soft-tissue resonance which occurs at heel-strike during walking and running. If this concept were true then the greatest vibration damping would occur when the input force was closest to the resonant frequency of the soft-tissues at heel-strike. However, this idea has not been tested. The purpose of this study was to test whether muscle activity in the lower extremity is used to damp soft-tissue resonance which occurs at heel-strike during walking. Hard and soft shoe conditions were tested in a randomized block design. Ground reaction forces, soft-tissue accelerations and myoelectric activity were measured during walking for 40 subjects. Soft-tissue mass was estimated from anthropologic measurements, allowing inertial forces in the soft-tissues to be calculated. The force transfer from the ground to the tissues was compared with changes in the muscle activity. The soft condition resulted in relative frequencies (input/tissue) to be closer to resonance for the main soft-tissue groups. However, no increase in force transmission was observed. Therefore, the vibration damping in the tissues must have increased. This increase concurred with increases in the muscle activity for the biceps femoris and lateral gastrocnemius. The evidence supports the proposal that muscle activity damps soft-tissue resonance at heel-strike. Muscles generate forces which act across the joints and, therefore, shoe design may be used to modify muscle activity and thus joint loading during walking and running.     

Masticatory function in patients with xerostomia

The effects of reduced salivary output in patients suffering from xerostomia on masticatory function has not been previously studied. This study compares masticatory performance and kinematic activity of patients suffering from xerostomia with age-, sex-, and number of occluding pairs-matched healthy controls. Masticatory function was evaluated by assessment of chewing motion and muscle activity during chewing an artificial food (CutterSil®), chewing gum and swallowing a bolus of almond. Chewing motion was recorded with the Optotrak® computer system. Bilateral muscle activity of both masseter and anterior temporalis was recorded using surface electrodes. Results of this study revealed significant differences between patients and controls in their ability to process food and masticatory muscle activity. The majority of patients could not break down the artificial food, others had a larger median particle size than the controls. A significant difference was also observed in the number of chewing cycles required to swallow almonds, the patients required more than twice as many chews as the controls, P<0.001. The right masseter muscle displayed significantly less activity for the patient than the controls. These findings suggest that patients with xerostomia exhibit reduced ability to process food. The observed decline in masticatory performance is probably due to reduced activity of the muscles of mastication.     

Muscle tuning during running: implications of an un-tuned landing

BACKGROUND: The impact force in heel-toe running is an input signal into the body that initiates vibrations of the soft tissue compartments of the leg. These vibrations are heavily damped and the paradigm of muscle tuning suggests the body adapts to different input signals to minimize these vibrations. The objectives of the present study were to investigate the implications of not tuning a muscle properly for a landing with a frequency close to the resonance frequency of a soft tissue compartment and to look at the effect of an unexpected surface change on the subsequent step of running. METHOD: Thirteen male runners were recruited and performed heel-toe running over two surface conditions. The peak accelerations and biodynamic responses of the soft tissue compartments of the leg along with the EMG activity of related muscles were determined for expected soft, unexpected hard and expected hard landings. RESULTS AND CONCLUSIONS: For the unexpected hard landing there was a change in the input frequency of the impact force, shifting it closer to the resonance frequency of the soft tissue compartments. For the unexpected landing there was no muscle adaptation, as subjects did not know the running surface was going to change. In support of the muscle-tuning concept an increase in the soft tissue acceleration did occur. This increase was greater when the proximity of the input signal frequency was closer to the resonance frequency of the soft tissue compartment. Following the unexpected change in the input signal a change in pre-contact muscle activity to minimize soft tissue compartment vibrations was not found. This suggests if muscle tuning does occur it is not a continuous feedback response that occurs with every small change in the landing surface properties. In previous studies with significant adaptation periods to new input signals significant correlations between the changes in the input signal frequency and the EMG intensity have been shown, however, changes in soft tissue accelerations have not been found. The results of the present study showed that changes in these soft tissue accelerations can occur in response to a resonance frequency input signal when a muscle reaction has not happened.     

Complexity of ruminant masticatory evolution

The evolution of robust jaws, hypsodont teeth, and large chewing muscles among grazing ruminants is a quintessential example of putative morphological adaptation. However, the degree of correlated evolution (i.e., to what extent the grazer feeding apparatus represents an evolutionary module), especially of soft and hard tissues, remains poorly understood. Recent generation of large datasets and phylogenetic information has made testing hypotheses of correlated evolution possible. We, therefore, test for correlated evolution among various traits of the ruminant masticatory apparatus including tooth crown height, jaw robustness, chewing muscle size, and characters of the molar occlusal surfaces, using phylogenetic and nonphylogenetic comparative methods as well as phylogenetic evolutionary model selection. We find that the large masseter muscles of grazing ruminants evolved with the inclusion of grass in the diet, an increase in the proportion of occlusal enamel bands oriented parallel to the chewing stroke, and possibly hypsodonty. We suggest that the masseter evolved under two evolutionary regimes: i) selection for higher masticatory forces during chewing and ii) flattening of the tooth profile, which resulted in reduced tooth guidance and, thus, a requirement for more chewing muscle activity during each chewing stroke, in agreement with previous research. The linear jaw metrics (depth of the mandibular angle, mandibular angle width, and length of the superficial masseteric scar) all show correlated evolution with hypsodonty and the proportion of enamel bands oriented parallel to the chewing stroke. We suggest that changes in the shape of the mandible represent the combined effects of selection for a reorientation of the chewing stroke, so as to emphasize horizontal translation of the teeth, and accommodation of high‐crowned teeth. Our analyses show that the ruminant feeding apparatus is an evolutionary mosaic with its various components showing both correlated and independent evolution. J. Morphol. 275:1093–1102, 2014. © 2014 Wiley Periodicals, Inc.     

Functional significance of the mandibular symphysis.

The purpose of this investigation was to relate the morphology of connective tissues in the mandibular symphysis to the behavioral and experimental evidence for mobility and mechanical stress at the symphysis. The anatomy of the symphysis was examined histologically in 6 mammalian orders encompassing 22 species. Behavioral and experimental evidence of stress during the power stroke of the chewing cycle correspond with stresses at the symphysis implied by the location and orientation of symphyseal connective tissues. These stresses are: (1) dorsoventral shear of the symphysis due to the transfer of force from balancing to chewing sides, (2) bending of the symphysis causing tension along the inferior and compression along superior borders due to torsion on the dentaries from the jaw closing muscles, and (3) antero-posterior shear of the symphysis due to an anteriorly directed stress on the chewing side. Interspecific comparisons suggest that leaf eaters can resist greater dorsoventral shear than fruit or insect eaters, but no correlations exist between diet and bending or antero-posterior shear. This suggests that chewing leaves requires larger biting forces.     

Temporally patterned activity recorded from mandibular nerves of the isolated subesophageal ganglion of Manduca

We recorded bursts of motor neuron activity from closer and opener mandibular nerves of isolated subesophageal ganglia (SOG) and compared them with the feeding motor pattern of intact Manduca larvae. Closer bursts recorded from isolated SOG lasted from 1 to 4s, interburst interval durations lasted from 2 to 49s, and within- and between-animal variability was great. In contrast, motor activity bursts (EMGs) measured from mandibular closer muscles of intact, feeding animals lasted 0.08 to 0.24s with interburst intervals of 0.26 to 0.57s. Variability both within and between animals was small. Bath application of 10(-4)M octopamine to the isolated SOG tended to increase frequency and reduce the duration of bursts, so that they became more like those recorded during feeding.     

Muscle activity damps the soft tissue resonance that occurs in response to pulsed and continuous vibrations.

This study tested the hypotheses that when the excitation frequency of mechanical stimuli to the foot was close to the natural frequency of the soft tissues of the lower extremity, the muscle activity increases 1) the natural frequency and 2) the damping to minimize resonance. Soft tissue vibrations were measured with triaxial accelerometers, and muscle activity was measured by using surface electromyography from the quadriceps, hamstrings, tibialis anterior, and triceps surae groups from 20 subjects. Subjects were presented vibrations while standing on a vibrating platform. Both continuous vibrations and pulsed bursts of vibrations were presented, across the frequency range of 10-65 Hz. Elevated muscle activity and increased damping of vibration power occurred when the frequency of the input was close to the natural frequency of each soft tissue. However, the natural frequency of the soft tissues did not change in a manner that correlated with the frequency of the input. It is suggested that soft tissue damping may be the mechanism by which resonance is minimized at heel strike during running.     

Task-specific recruitment of motor units for vibration damping

Vibrations occur within the soft tissues of the lower extremities due to the heel-strike impact during walking. Increases in muscle activity in the lower extremities result in increased damping to reduce this vibration. The myoelectric intensity spectra were compared using principal component analysis from the tibialis anterior and lateral gastrocnemius of 40 subjects walking with different shoe conditions. The soft insert condition resulted in a significant, simultaneous increase in muscle activity with a shift to higher myoelectric frequencies in the period 0-60 ms after heel-strike which is the period when the greater vibration damping occurred. These increases in myoelectric frequency match the spectral patterns which indicate increases in recruitment of faster motor units. It is concluded that fast motor units are recruited during the task of damping the soft-tissue resonance that occurs following heel-strike.     

Modification of soft tissue vibrations in the leg by muscular activity.

Vibration characteristics were recorded for the soft tissues of the triceps surae, tibialis anterior, and quadriceps muscles. The frequency and damping of free vibrations in these tissues were measured while isometric and isotonic contractions of the leg were performed. Soft tissue vibration frequency and damping increased with both the force produced by and the shortening velocity of the underlying muscle. Both frequency and damping were greater in a direction normal to the skin surface than in a direction parallel to the major axis of each leg segment. Vibration characteristics further changed with the muscle length and between the individuals tested. The range of the measured vibration frequencies coincided with typical frequencies of impact forces during running. However, observations suggest that soft tissue vibrations are minimal during running. These results support the strategy that increases in muscular activity may be used by some individuals to move the frequency and damping characteristics of the soft tissues away from those of the impact force and thus minimize vibrations during walking and running.     

Effect of bolus hardness on the chewing pattern and activation of masticatory muscles in subjects with normal dental occlusion

The aim of the study was to evaluate the effect of bolus hardness on the kinematic of mastication and jaw-elevator muscle activity in subjects with normal dental occlusion and function. The mandibular motion and the surface EMG envelope of the masseter and temporalis anterior muscles were assessed in twelve subjects during mastication of a soft and hard bolus of the same size. When chewing the hard bolus, the chewing pattern in the frontal plane was significantly higher and wider, with smaller closure angle and higher peak velocity than when chewing the soft bolus. EMG peak amplitude of both the masseter and anterior temporalis muscles was higher for the side of the bolus but the contralateral side increased its activity significantly more than the ipsilateral side when the hardness of the bolus increased (for the masseter, mean ± SD: 130.4 ± 108.1% increase for the contralateral side and 29.6 ± 26.9% for the ipsilateral side). Moreover, the peak EMG activity for both muscles occurred more distant from the closure point with hard bolus. The increased activity of the contralateral side may help maintaining the mandibular equilibrium, with indirect participation to the power stroke generated by the chewing-side masseter. The results provide kinematic and EMG adaptations to bolus hardness in healthy subjects and can be used as normative data in the development of methods for early diagnosis of impaired chewing function.     

Measurements of locomotor activity in hatchlings of the migratory locust Locusta migratoria: effects of intrinsic and extrinsic factors

Abstract. The effects of intrinsic and extrinsic factors on locomotor activity in crowd‐reared first‐stadium nymphs of the migratory locust Locusta migratoria are investigated under continuous light conditions using an actograph apparatus. Nymphs show monomodal or bimodal patterns of locomotor activity with respect to the time after the start of measurements, depending on the age. Locomotor activity is suppressed by the presence of grass in nymphs aged 0–2 days old but a peak of activity observed shortly after hatching is not suppressed. The results suggest that newly‐hatched nymphs may try to disperse from the hatching site. Nymphs show higher locomotor activity levels under moist conditions than under dry conditions during the first 5‐h period of measurements. This enhanced locomotor activity may constitute attempts to avoid high humidity. Under dim‐light conditions (2 × 10^?2 Wm^?2), locomotor activity is suppressed during the first half day and increases to a high level thereafter in both grass‐fed and unfed individuals. This increased activity might indicate a possible involvement of circadian rhythms. Background colour has no significant effect on the locomotor activity. The present study provides new aspects of behaviour in nymphs as well as baseline data for behavioural analysis of locust locomotion in relation to phase polyphenism.     

PYCNOGENOL chewing gum minimizes gingival bleeding and plaque formation.

PYCNOGENOL is an antioxidant phytochemical shown to have antiinflammatory activity in both the in vitro and in vivo models. This study compared the effects of chewing gums with and without PYCNOGENOL on gingival bleeding and plaque formation in 40 human subjects. In this double-blind study, subjects were assigned randomly to receive either control gums without PYCNOGENOL or experimental gums containng 5 mg PYCNOGENOL. Subjects used chewing gums for 14 days. Gingival bleeding and plaque scores were taken before and after the experiment. PYCNOGENOL chewing gums significantly reduced gingival bleeding, while no changes were noted in bleeding indexes in control subjects who used regular chewing gums. Subjects using regular control gums had significant increases of dental plaque accumulation during the two-week period. No increases in plaque accumulation were noted in subjects using PYCNOGENOL chewing gums. The data of this study suggest that the use of Pycnogenol chewing gums can minimize gingival bleeding and plaque accumulation.     

Manipulating field margins to increase predation intensity in fields of winter wheat (Triticum aestivum)

The effectiveness of natural enemies to control pests can be enhanced through habitat manipulation. However, due to the differences in their ecology, generalist and specialist species may respond differently to the same manipulation. Moreover, interactions among natural enemies (i.e. cannibalism, intraguild predation, hyperparasitism) may complicate the assumption that a higher density of natural enemies would increase the level of biological control. We investigated the natural enemy guild composition and the predation rate along flower vs. grass margins at the edge of winter wheat (Triticum aestivum) fields in Denmark. Natural enemies were sampled by pitfall trapping and by suction sampling; predation intensity was measured using two different sentinel prey methods: artificial caterpillars made of plasticine, and sentinel aphid colonies. Specialist and generalist species responded differently to the two margin types: specialists (mostly parasitic wasps) were attracted by the flower margins, while generalists (ground beetles, rove beetles and spiders) were more active in grass margins. The number of artificial caterpillars attacked was significantly greater in grass margins (mean = 48.9%, SD = 24.3) than in flower margins (mean = 30.7%, SD = 17.4). We found a significant positive relationship between the number of artificial caterpillars attacked by chewing insects, and activity density for large (≥15 mm) ground beetles. Predation of sentinel aphids in wheat fields did not vary significantly in relation to margin type. Our results suggest that flowering margins may be beneficial for canopy‐active specialist natural enemies, but grassy margins are more useful for ground‐active generalist predators.     

A new method of evaluation of the masticatory function in humans

A method is proposed to estimate mastication by three indices: chewing effect, chewing performance, chewing efficiency. Definitions are given for these terms. A new chewing test is described, permits studying the influence of the number of chewing strokes, chewing force and volume of test portion on the mastication using a group of 25 dentate persons aged 17-25 years with class I occlusion. A new approach to the estimation of chewing efficiency changes traditional views. A strong positive correlation is first found between chewing effect (A) and integrated bioelectrical activity (J) of chewing muscles with the constant number of chewing strokes. A relation between A and J is established when the number of chewing strokes increases.     

Feedback in the Contractile Mechanism of the Frog Heart

Shortening causes a transient decrease, extension an increase, in activity during contractures of the frog ventricle induced by high Ca or by isosmotic K solution. This is shown by the fact that, after the immediate passive shortening, the muscle is extended under isotonic conditions when the load is diminished, and that under isometric conditions quick release causes first a rapid drop, then a further, much slower, fall of tension. Increasing the load or stretching induce the opposite effects. At low temperatures all rapid changes in length produce oscillations of low frequency. These responses are due to a sensitive feedback mechanism similar to that previously demonstrated for insect fibrillar muscle. That this mechanism comes into play in the heart under normal conditions and controls the time-course of the twitch is demonstrated by the observation that relaxation begins earlier the greater the shortening. Thus, during afterloaded isotonic twitches the onset of relaxation is advanced as the load is diminished.     

Climatic influences on the behavioural ecology of Chanter's mountain reedbuck in Kenya

The effects of rainfall and temperature on the behavioural ecology of Chanler's mountain reedbuck (Redunca fulvorufula fulvorufula Rothschild) were examined on ranchland near Gilgil, Kenya. Ambient temperature was shown to be the proximate determinant of diurnal activity and rumination patterns. Mountain reedbuck were active during early morning and late afternoon, but rested and abandoned rumination when temperatures peaked at midday. There was close synchrony in levels of activity, rumination and use of cover and shade between males and females. Seasonal variations in time budgets were strongly influenced by rainfall patterns. Analyses revealed a one-month lag between rainfall and both peak grass growth and a decrease in rumination frequency. The proportion of time allocated to feeding decreased one month later, and was coincident with an increase in the proportion of grass in the diet. Reedbuck may therefore be prevented from exploiting high-quality new grass, possibly by gut-fill or induced imbalances in rumen pH. It is suggested that the unexpectedly high levels of browse in the diet is an adaptive response to low rainfall during the preceding two months.     

Chewing reduces sympathetic nervous response to stress and prevents poststress arrhythmias in rats

Reducing stress is important in preventing sudden death in patients with cardiovascular disease, as stressful events may cause autonomic imbalance and trigger fatal arrhythmias. Since chewing has been shown to inhibit stress-induced neuronal responses in the hypothalamus, we hypothesized that chewing could ameliorate stress-induced autonomic imbalance and prevent arrhythmias. To test this hypothesis, we analyzed changes in radiotelemetered electrocardiograms in rats that were allowed to chew a wooden stick during a 1-h period of immobilization stress. Chewing significantly reduced the occurrence of ventricular premature beats (VPBs) and complex ventricular ectopy after immobilization and prevented stress-induced prolongation of the QT interval of VPBs throughout the 10-h experimental period. It also prevented prolongation of the QRS complex and fluctuations in the QT interval in normal sinus rhythm beats preceding VPBs during both immobilization and in the poststress period. Fast Fourier transform-based spectral analysis of heart-rate variability further showed that chewing significantly inhibited the stress-induced increase in the power ratio of low-to-high frequency activity (LF/HF: a marker of sympathetic activity) during immobilization and in addition was associated with blunting of the stress-induced increase in plasma noradrenaline observed at the termination of immobilization. Similar suppressive effects on the occurrence of VPBs and the LF/HF were observed in rats that were administered the β-adrenergic blocker propranolol before immobilization. These results indicate that chewing can ameliorate sympathetic hyperactivity during stress and prevent poststress arrhythmias and suggest that chewing may provide a nonpharmacological and cost-effective treatment option for patients with a high risk of stress-induced fatal arrhythmia.