Circadian rhythms of heart rate in freely moving and restrained American lobsters, Homarus americanus

While circadian rhythms of locomotion have been reported in the American lobster, Homarus americanus, it is unclear whether heart rate is also modulated on a circadian basis. To address this issue, both heart rate and locomotor activity were continuously monitored in light-dark (LD) cycles and constant darkness (DD). Lobsters in running wheels exhibited significant nocturnal increases in locomotor activity and heart rates during LD, and these measures were significantly correlated. In DD, most lobsters exhibited persistent circadian rhythms of both locomotion and heart rate. When heart rate was monitored in restrained lobsters in LD and DD, most animals also demonstrated clear daily and circadian rhythms in heart rate. Overall, this is the first demonstration of circadian rhythms of heart rate in H. americanus, the expression of which does not appear to be dependent on the expression of locomotor activity.     

Juvenile hormone and circadian locomotor activity in the honey bee Apis mellifera

Age-related division of labor in honeybees is associated with plasticity in circadian rhythms. Young nest bees care for brood around the clock with no circadian rhythms while older foragers have strong circadian rhythms that are used for sun compass navigation and for timing visits to flowers. Since juvenile hormone (JH) is involved in the coordination of physiological and behavioral processes underlying age-related division of labor in honey bees, we tested the hypothesis that JH influences the ontogeny of circadian rhythms and other clock parameters in young worker bees. Treatments with the JH analog methoprene or allatectomy did not influence the onset of rhythmicity, overall locomotor activity, or the free-running period of rhythmic locomotor behavior. There were, however, significant differences in the onset of rhythmicity, overall locomotor activity, and longevity between bees from different source colonies, suggesting that there is significant genetic variation for these traits. Our results suggest that JH does not coordinate all aspects of division of labor in bees and that coordination of task performance with circadian rhythms is probably mediated by other regulatory systems.     

Circadian rhythms of heart rate in freely moving and restrained American lobsters,Homarus americanus

While circadian rhythms of locomotion have been reported in the American lobster, Homarus americanus, it is unclear whether heart rate is also modulated on a circadian basis. To address this issue, both heart rate and locomotor activity were continuously monitored in light-dark (LD) cycles and constant darkness (DD). Lobsters in running wheels exhibited significant nocturnal increases in locomotor activity and heart rates during LD, and these measures were significantly correlated. In DD, most lobsters exhibited persistent circadian rhythms of both locomotion and heart rate. When heart rate was monitored in restrained lobsters in LD and DD, most animals also demonstrated clear daily and circadian rhythms in heart rate. Overall, this is the first demonstration of circadian rhythms of heart rate in H. americanus, the expression of which does not appear to be dependent on the expression of locomotor activity.     

Circadian organization inAplysia: internal desynchronization and amplitude of locomotor rhythm

Summary We have tested the hypothesis that the circadian oscillators in the eyes ofAplysia are coequal driver oscillators for the circadian locomotor rhythm. Three predictions based on this hypothesis were tested. Prediction 1: at a time when the phase difference between the eye rhythms is small, the amplitude of the locomotor rhythm in two eyed animals will be as great or greater than the amplitude in one eyed animals. Prediction 2: the amplitude of the locomotor rhythm of two eyed animals will decline under conditions in which the two eye rhythms become out of phase with each other. Prediction 3: the form of the locomotor rhythm will broaden or become biphasic in two eyed animals when the two eye rhythms become out of phase with each other.None of the predictions was confirmed. One eyedAplysia had higher amplitude locomotor rhythms than two eyedAplysia, even under conditions in which the two eye rhythms were probably not far out of phase with each other. There was no tendency for the amplitude of the locomotor rhythm of two eyed animals to decline under circumstances in which the phase difference between the two eye rhythms changes from less than 4 h to as much as 11.5 h. There was no tendency in two eyed animals for the locomotor rhythm to broaden or become biphasic as the eye rhythms became more out of phase with each other.The results led us to reject the hypothesis that the eyes are co-equal drivers for the locomotor rhythm. The ocular influence on locomotion is more likely to be mediated via mechanisms in the central nervous system that do not faithfully conserve the phase of the eye rhythms. One possibility is that the driver is a third circadian oscillator that interacts with the two eye oscillators.Abbreviations CAP compound action potentials - CC constant conditions - CT circadian time - DO driver oscillator - EO eye oscillator - RSD relative standard deviations (see Methods)     

Rhythms of locomotion expressed by Limulus polyphemus, the American horseshoe crab: II. Relationship to circadian rhythms of visual sensitivity

In the laboratory, horseshoe crabs express a circadian rhythm of visual sensitivity as well as daily and circatidal rhythms of locomotion. The major goal of this investigation was to determine whether the circadian clock underlying changes in visual sensitivity also modulates locomotion. To address this question, we developed a method for simultaneously recording changes in visual sensitivity and locomotion. Although every animal (24) expressed consistent circadian rhythms of visual sensitivity, rhythms of locomotion were more variable: 44% expressed a tidal rhythm, 28% were most active at night, and the rest lacked statistically significant rhythms. When exposed to artificial tides, 8 of 16 animals expressed circatidal rhythms of locomotion that continued after tidal cycles were stopped. However, rhythms of visual sensitivity remained stable and showed no tendency to be influenced by the imposed tides or locomotor activity. These results indicate that horseshoe crabs possess at least two biological clocks: one circadian clock primarily used for modulating visual sensitivity, and one or more clocks that control patterns of locomotion. This arrangement allows horseshoe crabs to see quite well while mating during both daytime and nighttime high tides.     

Coupling between respiratory and stepping rhythms during locomotion in decerebrate cats

To determine whether and how the strength of coupling between respiratory and stepping rhythms varies depending on locomotor patterns, correlation analysis was done of diaphragmatic and gastrocnemius muscle activities. In spontaneously breathing cats decerebrated at the precollicular-post-mammillary level, tonic electrical stimulation was delivered to the mesencephalic locomotor region to induce locomotion on a treadmill. Electromyograms were recorded from the left hemidiaphragm and the bilateral gastrocnemius muscles. Various locomotor patterns were elicited by changes in the belt speed of the treadmill and in the intensity of stimulation of the mesencephalic locomotor region. Cross-correlograms between diaphragmatic and gastrocnemius activities showed that coupling was absent or weak when the cats walked slowly. The strength of locomotor-respiratory coupling tended to increase as the mean stepping interval shortened. When the animals were galloping, the respiratory rhythm was entrained 1:1 with the stepping rhythm. This study showed that the strength of coupling between respiratory and stepping rhythms varied depending on the locomotor patterns elicited, especially on whether the animals were running.     

Thoracic Hemisection in Rats Results in Initial Recovery Followed by a Late Decrement in Locomotor Movements,with Changes in Coordination Correlated with Serotonergic Innervation of the Ventral Horn

Lateral thoracic hemisection of the rodent spinal cord is a popular model of spinal cord injury, in which the effects of various treatments, designed to encourage locomotor recovery, are tested. Nevertheless, there are still inconsistencies in the literature concerning the details of spontaneous locomotor recovery after such lesions, and there is a lack of data concerning the quality of locomotion over a long time span after the lesion. In this study, we aimed to address some of these issues. In our experiments, locomotor recovery was assessed using EMG and CatWalk recordings and analysis. Our results showed that after hemisection there was paralysis in both hindlimbs, followed by a substantial recovery of locomotor movements, but even at the peak of recovery, which occurred about 4 weeks after the lesion, some deficits of locomotion remained present. The parameters that were abnormal included abduction, interlimb coordination and speed of locomotion. Locomotor performance was stable for several weeks, but about 3–4 months after hemisection secondary locomotor impairment was observed with changes in parameters, such as speed of locomotion, interlimb coordination, base of hindlimb support, hindlimb abduction and relative foot print distance. Histological analysis of serotonergic innervation at the lumbar ventral horn below hemisection revealed a limited restoration of serotonergic fibers on the ipsilateral side of the spinal cord, while on the contralateral side of the spinal cord it returned to normal. In addition, the length of these fibers on both sides of the spinal cord correlated with inter- and intralimb coordination. In contrast to data reported in the literature, our results show there is not full locomotor recovery after spinal cord hemisection. Secondary deterioration of certain locomotor functions occurs with time in hemisected rats, and locomotor recovery appears partly associated with reinnervation of spinal circuitry by serotonergic fibers.     

Coordination of wingbeat and respiration in birds. II. "Fictive" flight.

To determine whether an interaction between central respiratory and locomotor networks may be involved in the observed coordination of wingbeat and respiratory rhythms during free flight in birds, we examined the relationship between wingbeat and respiratory activity in decerebrate Canada geese and Pekin ducks before and after paralysis. Locomotor activity was induced through electrical stimulation of brain stem locomotor regions. Respiratory frequency (fv) was monitored via pneumotachography and intercostal electromyogram recordings before paralysis and via intercostal and cranial nerve IX electroneurogram recordings after paralysis. Wingbeat frequency (fW) was monitored using pectoralis major electromyogram recordings before, and electroneurogram recordings after, paralysis. Respiratory and cardiovascular responses of decerebrate birds during active (nonparalyzed) and "fictive" (paralyzed) wing activity were qualitatively similar to those of a variety of vertebrate species to exercise. As seen during free flight, wingbeat and respiratory rhythms were always coordinated during electrically induced wing activity. Before paralysis during active wing flapping, coupling ratios (fW/fv) of 1:1, 2:1, 3:1, and 4:1 (wingbeats per breath) were observed. After paralysis, fW and fv remained coupled; however, 1:1 coordination predominated. All animals tested (n = 9) showed 1:1 coordination. Two animals also showed brief periods of 2:1 coupling. It is clear that locomotor and respiratory networks interact on a central level to produce a synchronized output. The observation that the coordination between fW and fv differs in paralyzed and nonparalyzed birds suggests that peripheral feedback is involved in the modulation of a centrally derived coordination.     

Coordination of locomotor activity of mollusc Lymnaea stagnalis at nutrition: Role of the internal medium acid-base balance (pH)

The correlation was studied between the hemolymph pH value and the locomotion rate in the mollusc Lymnaea stagnalis at different degree of activation of the food-procuring behavior. It is established that the hemolymph pH value remains unchanged in the satiated (hungry), freely moving Lymnaea individuals, and, hence, it does not determine the locomotor activity of molluscs in these functional states. In the course of food consumption, locomotion decreases significantly, which is due to an observed decrease of the hemolymph pH. This effect is realized through the corresponding changes of electrical activity of motoneurons of the locomotor network of the Lymnaea nervous system (PeA-cluster). It is suggested that the pH value plays the modulating role in coordination of activities of the nervous centers of the mollusc nutrition and locomotion.     

Circadian locomotor rhythm masked by the female reproduction cycle in cockroaches

Blattella bisignata (Brunner) and B. germanica (L.) are oviparous cockroaches with cyclic reproductive behaviour, but in B. germanica only males show circadian rhythmicity of locomotion at 28°C and DD (constant darkness). In B. bisignata, males and virgin females cockroaches entrained by light–dark cycles show free‐running rhythmicity in DD, and most activities occur during the subjective night. Daily locomotor activities of virgin females show cyclic changes that coincided with ovarian development. Virgin females also exhibit calling behaviour during the subjective night, and this shows a free‐running rhythm. Male mate‐finding locomotion and female calling behaviour are under circadian control, so the timing for both behaviours is synchronized. However, most mated females do not show a locomotor free‐running rhythm under DD conditions. Our results indicate that only mated females could not express a circadian locomotor rhythm. Pregnancy reduces a female’s locomotory intensity and masks the expression of a circadian locomotor rhythm. We attribute the differences in circadian locomotory rhythms between these two species to their living environments and mate‐finding strategies.     

Intramuscular pressure-induced inhibition of cardiac contraction: implications for cardiac-locomotor synchronization

The synchronization of cardiac and locomotor rhythms has been suggested to enhance the efficiency of arterial delivery to active muscles during rhythmic exercise, but direct evidence showing such a functional role has not been provided. In this study, we tested the hypothesis that the heartbeat is coupled with intramuscular pressure (IMP) changes so as to time the delivery of blood through peripheral tissues when the IMP is lower. To this end, we developed a computer-controlled, dynamic, thigh cuff occlusion device that enables bilateral thigh cuffs to repeatedly inflate and deflate, one side after the other, to simulate rhythmic IMP changes during bipedal locomotion. Nine healthy subjects were examined, and three different occlusion pressures (50, 80, and 120 mmHg) were applied separately to the thigh cuffs of normal subjects while they were sitting. Alternate occlusions of the bilateral thigh cuffs administered at the frequency of the mean heart rate produced significant phase synchronization between the cardiac and cuff-occlusion rhythms when 120 mmHg pressure was applied. However, synchronization was not observed when the occlusion pressure was 50 or 80 mmHg. During synchronization, heartbeats were most likely to occur in phases that did not include overlap between the peak arterial flow velocity in the thigh and elevated cuff pressure. We believe that phase synchronization occurs so that the cardiac cycle is timed to deliver blood through the lower legs when IMP is not maximal. If this can be extrapolated to natural locomotion, synchronization between cardiac and locomotor activities may be associated with the improved perfusion of exercising muscles.     

Interactions between locomotion and ventilation in tetrapods

Interactions between locomotion and ventilation have now been studied in several species of reptiles, birds and mammals, from a variety of perspectives. Among these perspectives are neural interactions of separate but linked central controllers; mechanical impacts of locomotion upon ventilatory pressures and flows; and the extent to which the latter may affect gas exchange and the energetics of exercise. A synchrony, i.e. 1:1 pattern of coordination, is observed in many running mammals once they achieve galloping speeds, as well as in flying bats, some flying birds and hopping marsupials. Other, non-1:1, patterns of coordination are seen in trotting and walking quadrupeds, as well as running bipedal humans and running and flying birds. There is evidence for an energetic advantage to coordination of locomotor and respiratory cycles for flying birds and running mammals. There is evidence for a mechanical constraint upon ventilation by locomotion for some reptiles (e.g. iguana), but not for others (e.g. varanids and crocodilians). In diving birds the impact of wing flapping or foot paddling on differential air sac pressures enhances gas exchange during the breath hold by improving diffusive and convective movement of air sac oxygen to parabronchi. This paper will review the current state of our knowledge of such influences of locomotion upon respiratory system function.     

Interlimb coordination,gait, and neural control of quadrupedalism in chimpanzees

Interlimb coordination is directly relevant to the understanding of the neural control of locomotion, but few studies addressing this topic for nonhuman primates are available, and no data exist for any hominoid other than humans. As a follow-up to Jungers and Anapol's ([1985] Am. J. Phys. Anthropol. 67:89–97) analysis on a lemur and talapoin monkey, we describe here the patterns of interlimb coordination in two chimpanzees as revealed by electromyography. Like the lemur and talapoin monkey, ipsilateral limb coupling in chimpanzees is characterized by variability about preferred modes within individual gaits. During symmetrical gaits, limb coupling patterns in the chimpanzee are also influenced by kinematic differences in hindlimb placement (“overstriding”). These observations reflect the neurological constraints placed on locomotion but also emphasize the overall flexibility of locomotor neural mechanisms. Interlimb coordination patterns are also species-specific, exhibiting significant differences among primate taxa and between primates and cats. Interspecific differences may be suggestive of phylogenetic divergence in the basic mechanisms for neural control of locomotion, but do not preclude morphological explanations for observed differences in interlimb coordination across species. Am J Phys Anthropol 102:177–186, 1997 © 1997 Wiley-Liss, Inc.     

Development and relationships of locomotor, feeding, and oxygen consumption rhythms in house crickets

ABSTRACT. Locomotor, feeding, drinking, and oxygen consumption rhythms in adult virgin Acheta domesticus L. all appear to peak in the first half of the scotophase, be entrained cophasically by a LD 14:10 h cycle, have a lights-off Zeitgeber and persist in LL with a π c. 25 h for the locomotor rhythm and c. 23 h for the oxygen consumption rhythm. There is no evidence of these rhythms in last instar larvae. The onset of the locomotor rhythm requires 3 days at 30°C but 5–7 days at 25–28°C after the final ecdysis in virgins, indicating a temperature related development of the locomotor rhythm. Oxygen consumption rhythms are lacking in 2-day-old virgins but present in 8-day-old virgins. Feeding rhythms can be recorded in virgins as young as 2 days (before locomotor rhythm developed). Both oxygen consumption and locomotor rhythms persist during starvation. The results suggest that a central brain oscillator drives both feeding and locomotor rhythms independently, but that the oxygen consumption rhythm is derived from the metabolic demands associated with the other rhythms.     

Circadian rhythms of locomotor activity and temperature selection in sleepy lizards, <Emphasis Type="Italic">Tiliqua rugosa</Emphasis>

This study examined whether the daily rhythms of locomotor activity and behavioural thermoregulation that have previously been observed in Australian sleepy lizards (Tiliqua rugosa) under field conditions are true circadian rhythms that persist in constant darkness (DD) and whether these rhythms show similar characteristics. Lizards held on laboratory thermal gradients in the Australian spring under the prevailing 12-hour light : dark (LD) cycle for 14 days displayed robust daily rhythms of behavioural thermoregulation and locomotor activity. In the 13-day period of DD that followed LD, most lizards exhibited free-running circadian rhythms of locomotor activity and behavioural thermoregulation. The predominant activity pattern displayed in LD was unimodal and this was retained in DD. While mean levels of skin temperature and locomotor activity were found to decrease from LD to DD, activity duration remained unchanged. The present results demonstrate for the first time that this species’ daily rhythm of locomotor activity is an endogenous circadian rhythm. Our results also demonstrate a close correlation between the circadian activity and thermoregulatory rhythms in this species indicating that the two rhythms are controlled by the same master oscillator(s). Future examination of seasonal aspects of these rhythms, may, however, cause this hypothesis to be modified.     

Increased size due to larval royal jelly exposure does not affect circadian locomotor activity or climbing ability in adult female Drosophila melanogaster

The effects of royal jelly (RJ) appear to be conserved in Drosophila; flies exposed of RJ exhibit increased body size, similar to queen bees. However, in flies and bees, there is evidence that increased body size can lead to impairments to locomotor activity, while RJ may have anti-fatigue properties. Canton-S and Oregon-R Drosophila larvae were raised on media containing 0% or 20% pure RJ. Climbing assays were conducted to assess vertical locomotion. Circadian locomotion was observed using Drosophila Activity Monitors. CS, but not Or-R, raised on RJ were larger compared to controls. Flies exposed to RJ exhibited entrainment and free-running rhythms. The increased size due to RJ exposure in this study had no bearing on circadian locomotor activity or climbing. These results indicate that there is variation among physiological responses to RJ among different strains, but RJ was equally ineffective in affecting locomotor behavior no matter the physiological response.     

Disruption of Daily Rhythms by High-Fat Diet Is Reversible

In mammals a network of circadian clocks coordinates behavior and physiology with 24-h environmental cycles. Consumption of high-fat diet disrupts this temporal coordination by advancing the phase of the liver molecular clock and altering daily rhythms of eating behavior and locomotor activity. In this study we sought to determine whether these effects of high-fat diet on circadian rhythms were reversible. We chronically fed mice high-fat diet and then returned them to low-fat chow diet. We found that the phase of the liver PERIOD2::LUCIFERASE rhythm was advanced (by 4h) and the daily rhythms of eating behavior and locomotor activity were altered for the duration of chronic high-fat diet feeding. Upon diet reversal, the eating behavior rhythm was rapidly reversed (within 2 days) and the phase of the liver clock was restored by 7 days of diet reversal. In contrast, the daily pattern of locomotor activity was not restored even after 2 weeks of diet reversal. Thus, while the circadian system is sensitive to changes in the macronutrient composition of food, the eating behavior rhythm and liver circadian clock are specifically tuned to respond to changes in diet.     

Bio-inspired design strategies for central pattern generator control in modular robotics

New findings in the nervous system of invertebrates have shown how a number of features of central pattern generator (CPG) circuits contribute to the generation of robust flexible rhythms. In this paper we consider recently revealed strategies that living CPGs follow to design CPG control paradigms for modular robots. To illustrate them, we divide the task of designing an example CPG for a modular robot into independent problems. We formulate each problem in a general way and provide a bio-inspired solution for each of them: locomotion information coding, individual module control and inter-module coordination. We analyse the stability of the CPG numerically, and then test it on a real robot. We analyse steady state locomotion and recovery after perturbations. In both cases, the robot is able to autonomously find a stable effective locomotion state. Finally, we discuss how these strategies can result in a more general design approach for CPG-based locomotion.     

Drosophila olfactory response rhythms require clock genes but not pigment dispersing factor or lateral neurons

Odors elicit a number of behavioral responses, including attraction and repulsion in Drosophila. In this study, the authors used a T-maze apparatus to show that wild-type Drosophila melanogaster exhibit a robust circadian rhythm in the olfactory attractive and repulsive responses. These responses were lower during the day and began to rise at early night, peaking at about the middle of the night and then declining thereafter. They were also independent of locomotor activity. The olfactory response rhythms were lost in period or timeless mutant flies (per0, tim0), indicating that clock genes control circadian rhythms of olfactory behavior. The rhythms in olfactory response persisted in the absence of the pigment-dispersing factor neuropeptide or the central pacemaker lateral neurons known to drive circadian patterns of locomotion and eclosion. These results indicate that the circadian rhythms in olfactory behavior in Drosophila are driven by pacemakers that do not control the rest-activity cycle and are likely in the antennae.