Feeding Motor Patterns in Anurans: Insights from Biomechanical Modeling

During feeding in anurans, the mouth opens while the tongue,which is attached to the mandible at the front of the mouth,rotates forward. Due to the relative simplicity of its anatomyand the complexity of its motion, tongue protraction in frogspresents an ideal system for exploring the neural control ofmultijoint movements. In this study, we used a forward dynamic,rigid body model with four segments and two muscles to investigateopen loop control of tongue protraction in the Australian white-lippedtree frog, Litoria caerulea. Model parameters include the massdistribution, initial position and initial angular velocityof each segment and the anatomy and physiology of each muscle.Model variables include the level of muscle activation at eachtime step and impulsive torques to open and close the mouth.The model gives X,Y coordinates of each segment and joint anglesat each time step as output. The model was tested using scaled,normalized EMG signals and impulsive joint torques to predictthe paths of the lower jaw tip and tongue tip. Predicted pathswere compared to experimentally observed paths using Pearsonproduct-moment correlation coefficients. Simulations demonstratethat the genioglossus muscles likely play a minor role, if any,in determining the trajectory of the tongue in most anurans.Most of the force for tongue protraction comes from angularmomentum transferred to the tongue by the opening jaws. In anurans,tongue protraction is dynamically stable and will occur as longas the musculoskeletal elements are in the correct initial position.     

Localization of Motor Neurons and Central Pattern Generators for Motor Patterns Underlying Feeding Behavior in Drosophila Larvae

Motor systems can be functionally organized into effector organs (muscles and glands), the motor neurons, central pattern generators (CPG) and higher control centers of the brain. Using genetic and electrophysiological methods, we have begun to deconstruct the motor system driving Drosophila larval feeding behavior into its component parts. In this paper, we identify distinct clusters of motor neurons that execute head tilting, mouth hook movements, and pharyngeal pumping during larval feeding. This basic anatomical scaffold enabled the use of calcium-imaging to monitor the neural activity of motor neurons within the central nervous system (CNS) that drive food intake. Simultaneous nerve- and muscle-recordings demonstrate that the motor neurons innervate the cibarial dilator musculature (CDM) ipsi- and contra-laterally. By classical lesion experiments we localize a set of CPGs generating the neuronal pattern underlying feeding movements to the subesophageal zone (SEZ). Lesioning of higher brain centers decelerated all feeding-related motor patterns, whereas lesioning of ventral nerve cord (VNC) only affected the motor rhythm underlying pharyngeal pumping. These findings provide a basis for progressing upstream of the motor neurons to identify higher regulatory components of the feeding motor system.     

The Evolution of Feeding Motor Patterns in Lizards: Modulatory Complexity and Possible Constraints

Previous research indicated that the evolution of feeding motorpatterns across major taxonomic groups might have occurred withoutlarge modifications of the control of the jaw and hyolingualmuscles. However, the proposal of this evolutionary scheme washampered by the lack of data for some key taxa such as lizards.Recent data on jaw and hyolingual feeding motor patterns ofa number of lizard families suggest extensive variability withinand among species. Although most lizards respond to changesin the structural properties of food items by modulating theactivation of the jaw and hyolingual muscles, some food specialistsmight have lost this ability. Whereas the overall similarityin motor patterns across different lineages of lizards is largefor the hyolingual muscles, jaw muscle activation patterns seemto be more flexible. Nevertheless, all data suggest that boththe jaw and hyolingual system are complexly integrated. Theelimination of feedback pathways from the hyolingual systemthrough nerve transection experiments clearly shows that feedingcycles are largely shaped by feedback interactions. Yet, novelmotor patterns including unilateral control seem to have emergedin the evolution from lizards to snakes.     

Patterns of positive selection in six Mammalian genomes

Genome-wide scans for positively selected genes (PSGs) in mammals have provided insight into the dynamics of genome evolution, the genetic basis of differences between species, and the functions of individual genes. However, previous scans have been limited in power and accuracy owing to small numbers of available genomes. Here we present the most comprehensive examination of mammalian PSGs to date, using the six high-coverage genome assemblies now available for eutherian mammals. The increased phylogenetic depth of this dataset results in substantially improved statistical power, and permits several new lineage- and clade-specific tests to be applied. Of approximately 16,500 human genes with high-confidence orthologs in at least two other species, 400 genes showed significant evidence of positive selection (FDR<0.05), according to a standard likelihood ratio test. An additional 144 genes showed evidence of positive selection on particular lineages or clades. As in previous studies, the identified PSGs were enriched for roles in defense/immunity, chemosensory perception, and reproduction, but enrichments were also evident for more specific functions, such as complement-mediated immunity and taste perception. Several pathways were strongly enriched for PSGs, suggesting possible co-evolution of interacting genes. A novel Bayesian analysis of the possible "selection histories" of each gene indicated that most PSGs have switched multiple times between positive selection and nonselection, suggesting that positive selection is often episodic. A detailed analysis of Affymetrix exon array data indicated that PSGs are expressed at significantly lower levels, and in a more tissue-specific manner, than non-PSGs. Genes that are specifically expressed in the spleen, testes, liver, and breast are significantly enriched for PSGs, but no evidence was found for an enrichment for PSGs among brain-specific genes. This study provides additional evidence for widespread positive selection in mammalian evolution and new genome-wide insights into the functional implications of positive selection.     

Characteristic Species Dependent Growth Patterns of Mammalian Neoplasms

Evidence is presented, arising from an analysis of published data on tumour growth in three species of laboratory animals and in human multiple myeloma supporting a species specific relation between two supposedly independent parameters in the Gompertz equations frequently used to quantify tumour growth curves. This evidence supports the conjecture of Norton et al. (1976), based on their observations of the growth kinetics of a murine melanoma and a rat mammary carcinoma, that such a relation may be a general feature of tumour growth. Published data on the growth of xenografts of human colorectal tumours in immune-deprived mice suggests that the observed growth relation reflects the ability of a particular species to support a tumour of a certain maximum size. the existence of this relation greatly simplifies the task of predicting complete patterns of undisturbed neoplastic growth in these species.     

Patterns of Insertion and Deletion in Mammalian Genomes

Nucleotide insertions and deletions (indels) are responsible for gaps in the sequence alignments. Indel is one of the major sources of evolutionary change at the molecular level. We have examined the patterns of insertions and deletions in the 19 mammalian genomes, and found that deletion events are more common than insertions in the mammalian genomes. Both the number of insertions and deletions decrease rapidly when the gap length increases and single nucleotide indel is the most frequent in all indel events. The frequencies of both insertions and deletions can be described well by power law.Key Words: Insertion, deletion, gap, indel, mammalian genome.     

Mammalian Rest/Activity Patterns Explained by Physiologically Based Modeling

Circadian rhythms are fundamental to life. In mammals, these rhythms are generated by pacemaker neurons in the suprachiasmatic nucleus (SCN) of the hypothalamus. The SCN is remarkably consistent in structure and function between species, yet mammalian rest/activity patterns are extremely diverse, including diurnal, nocturnal, and crepuscular behaviors. Two mechanisms have been proposed to account for this diversity: (i) modulation of SCN output by downstream nuclei, and (ii) direct effects of light on activity. These two mechanisms are difficult to disentangle experimentally and their respective roles remain unknown. To address this, we developed a computational model to simulate the two mechanisms and their influence on temporal niche. In our model, SCN output is relayed via the subparaventricular zone (SPZ) to the dorsomedial hypothalamus (DMH), and thence to ventrolateral preoptic nuclei (VLPO) and lateral hypothalamus (LHA). Using this model, we generated rich phenotypes that closely resemble experimental data. Modulation of SCN output at the SPZ was found to generate a full spectrum of diurnal-to-nocturnal phenotypes. Intriguingly, we also uncovered a novel mechanism for crepuscular behavior: if DMH/VLPO and DMH/LHA projections act cooperatively, daily activity is unimodal, but if they act competitively, activity can become bimodal. In addition, we successfully reproduced diurnal/nocturnal switching in the rodent Octodon degu using coordinated inversions in both masking and circadian modulation. Finally, the model correctly predicted the SCN lesion phenotype in squirrel monkeys: loss of circadian rhythmicity and emergence of ∼4-h sleep/wake cycles. In capturing these diverse phenotypes, the model provides a powerful new framework for understanding rest/activity patterns and relating them to underlying physiology. Given the ubiquitous effects of temporal organization on all aspects of animal behavior and physiology, this study sheds light on the physiological changes required to orchestrate adaptation to various temporal niches.     

A Neural Infrastructure for Rhythmic Motor Patterns

It is possible to work out the neural circuity of many invertebrate central pattern generators (CPGs) thereby providing a basis for linking cellular processes to actual behaviors. This review summarizes the infrastructure of the two CPGs in the lobster stomatogastric ganglion in terms of circuitry, ionic conductances and chemical modulation by amines and peptides. Analysis of the circuit using modeling techniques including the use of electronic neurons closes the chapter.     

Motor Training Increases the Stability of Activation Patterns in the Primary Motor Cortex

Learning to be skillful is an endowed talent of humans, but neural mechanisms underlying behavioral improvement remain largely unknown. Some studies have reported that the mean magnitude of neural activation is increased after learning, whereas others have instead shown decreased activation. In this study, we used functional magnetic resonance imaging (fMRI) to investigate learning-induced changes in the neural activation in the human brain with a classic motor training task. Specifically, instead of comparing the mean magnitudes of activation before and after training, we analyzed the learning-induced changes in multi-voxel spatial patterns of neural activation. We observed that the stability of the activation patterns, or the similarity of the activation patterns between the even and odd runs of the fMRI scans, was significantly increased in the primary motor cortex (M1) after training. By contrast, the mean magnitude of neural activation remained unchanged. Therefore, our study suggests that learning shapes the brain by increasing the stability of the activation patterns, therefore providing a new perspective in understanding the neural mechanisms underlying learning.     

Prey Capture in Actinopterygian Fishes: A Review of Suction Feeding Motor Patterns with New Evidence from an Elopomorph Fish, Megalops atlanticus

Suction feeding is recognized as the dominant mode of aquaticprey capture in fishes. While much work has been done identifyingmotor pattern variations of this behavior among diverse groupsof actinopterygian fishes, many ray-finned groups are stillnot represented. Further, the substantial amount of inherentvariation in electromyography makes much of the pioneering workof suction feeding motor patterns in several basal groups insufficientfor evolutionary comparisons. Robust evolutionary comparisonshave identified conserved qualitative traits in the order ofmuscle activation during suction feeding (jaw opening > buccalcavity expansion > jaw closing). However, quantitative traitsof suction motor patterns (i.e., burst durations and relativeonset times) have changed over evolutionary time among actinopterygianfishes. Finally, new motor pattern evidence is presented froma previously neglected group, the Elopomorpha. The results suggestthat future investigations of the muscles influencing lateralexpansion of the mouth cavity and head anatomy may provide valuablenew insights into the evolution of suction feeding motor patternsin ray-finned fishes. In addition, the evidence illustratesthe value of comprehensive EMG surveys of cranial muscle activitiesduring suction feeding behavior.     

Patterns of Evolution in the Feeding Mechanism of Actinopterygian Fishes

SYNOPSIS. Structural and functional patterns in the evolutionof the actinopterygian feeding mechanism are discussed in thecontext of the major monophyletic lineages of ray-finned fishes.A tripartite adductor mandibulae contained in a maxillary-palatoquadratechamber and a single mechanism of mandibular depression mediatedby the obliquus inferioris, sternohyoideus, and hyoid apparatusare primitive features of the Actinopterygii. Halecostome fishesare characterized by having an additional mechanism of mandibulardepression, the levator operculi—opercular series coupling,and a maxilla which swings anteriorly during prey capture. Theseinnovations provide the basis for feeding by inertial suctionwhich is the dominant mode of prey capture throughout the halecostomeradiation. A remarkably consistent kinematic profile occursin all suction-feeding halecostomes. Teleost fishes possessa number of specializations in the front jaws including a geniohyoideusmuscle, loss of the primitive suborbital adductor component,and a mobile premaxilla. Structural innovations in teleost pharyngealjaws include fusion of the dermal tooth plates with endoskeletalgill arch elements, the occurrence of a pharyngeal retractormuscle, and a shift in the origin of the pharyngohyoideus. Thesespecializations relate to increased functional versatility ofthe pharyngeal jaw apparatus as demonstrated by an electromyographicstudy of pharyngeal muscle activity in Esox and Ambloplites.The major feature of the evolution of the actinopterygian feedingmechanism is the increase in structural complexity in both thepharyngeal and front jaws. Structural diversification is a functionof the number of independent biomechanical pathways governingmovement.     

The Evolution of the Motor Control of Feeding in Amphibians

Based on studies of a few model taxa, amphibians have been consideredstereotyped in their feeding movements relative to other vertebrates.However, recent studies on a wide variety of amphibian specieshave revealed great diversity in feeding mechanics and kinematics,and illustrate that stereotypy is the exception rather thanthe rule in amphibian feeding. Apparent stereotypy in some taxamay be an artifact of unnatural laboratory conditions. The commonancestor of lissamphibians was probably capable of some modulationof feeding movements, and descendants have evolved along twotrajectories with regard to motor control: (1) an increase inmodulation via feedback or feed-forward mechanisms, as exemplifiedby ballistic-tongued plethodontid salamanders and hydrostatic-tonguedfrogs, and (2) a decrease in variation dictated by biomechanicsthat require tight coordination between different body parts,such as the tongue and jaws in toads and other frogs with ballistictongue projection. Multi-joint coordination of rapid movementsmay hamper accurate tongue placement in ballistic-tongued frogsas compared to both short-tongued frogs and ballistic tongued-salamandersthat face simpler motor control tasks. Decoupling of tongueand jaw movements is associated with increased accuracy in bothhydrostatic-tongued frogs and ballistic-tongued salamanders.     

Stimulation of Glia Reveals Modulation of Mammalian Spinal Motor Networks by Adenosine

Despite considerable evidence that glia can release modulators to influence the excitability of neighbouring neurons, the importance of gliotransmission for the operation of neural networks and in shaping behaviour remains controversial. Here we characterise the contribution of glia to the modulation of the mammalian spinal central pattern generator for locomotion, the output of which is directly relatable to a defined behaviour. Glia were stimulated by specific activation of protease-activated receptor-1 (PAR1), an endogenous G-protein coupled receptor preferentially expressed by spinal glia during ongoing activity of the spinal central pattern generator for locomotion. Selective activation of PAR1 by the agonist TFLLR resulted in a reversible reduction in the frequency of locomotor-related bursting recorded from ventral roots of spinal cord preparations isolated from neonatal mice. In the presence of the gliotoxins methionine sulfoximine or fluoroacetate, TFLLR had no effect, confirming the specificity of PAR1 activation to glia. The modulation of burst frequency upon PAR1 activation was blocked by the non-selective adenosine-receptor antagonist theophylline and by the A₁-receptor antagonist 8-cyclopentyl-1,3-dipropylxanthine, but not by the A_2A-receptor antagonist SCH5826, indicating production of extracellular adenosine upon glial stimulation, followed by A₁-receptor mediated inhibition of neuronal activity. Modulation of network output following glial stimulation was also blocked by the ectonucleotidase inhibitor ARL67156, indicating glial release of ATP and its subsequent degradation to adenosine rather than direct release of adenosine. Glial stimulation had no effect on rhythmic activity recorded following blockade of inhibitory transmission, suggesting that glial cell-derived adenosine acts via inhibitory circuit components to modulate locomotor-related output. Finally, the modulation of network output by endogenous adenosine was found to scale with the frequency of network activity, implying activity-dependent release of adenosine. Together, these data indicate that glia play an active role in the modulation of mammalian locomotor networks, providing negative feedback control that may stabilise network activity.     

Feeding and Stocking Up: Radio-Labelled Food Reveals Exchange Patterns in Ants

Food sharing is vital for a large number of species, either solitary or social, and is of particular importance within highly integrated societies, such as in colonial organisms and in social insects. Nevertheless, the mechanisms that govern the distribution of food inside a complex organizational system remain unknown.Using scintigraphy, a method developed for medical imaging, we were able to describe the dynamics of food-flow inside an ant colony. We monitored the sharing process of a radio-labelled sucrose solution inside a nest of Formica fusca. Our results show that, from the very first load that enters the nest, food present within the colony acts as negative feedback to entering food. After one hour of the experiments, 70% of the final harvest has already entered the nest. The total foraged quantity is almost four times smaller than the expected storage capacity. A finer study of the spatial distribution of food shows that although all ants have been fed rapidly (within 30 minutes), a small area representing on average 8% of the radioactive surface holds more than 25% of the stored food. Even in rather homogeneous nests, we observed a strong concentration of food in few workers. Examining the position of these workers inside the nest, we found heavily loaded ants in the centre of the aggregate. The position of the centre of this high-intensity radioactive surface remained stable for the three consecutive hours of the experiments.We demonstrate that the colony simultaneously managed to rapidly feed all workers (200 ants fed within 30 minutes) and build up food stocks to prevent food shortage, something that occurs rather often in changing environments. Though we expected the colony to forage to its maximum capacity, the flow of food entering the colony is finely tuned to the colony''s needs. Indeed the food-flow decreases proportionally to the food that has already been harvested, liberating the work-force for other tasks.     

Catch Me If You Can: Diel Activity Patterns of Mammalian Prey and Predators

The activity patterns exhibited by animals are shaped by evolution, but additionally fine‐tuned by flexible responses to the environment. Predation risk and resource availability are environmental cues which influence the behavioural decisions that make both predators and prey engage in activity bursts, and depending on their local importance, can be strong enough to override the endogenous regulation of an animals’ circadian clock. In Southern Europe, wherever the European rabbit (Oryctolagus cuniculus) is abundant, it is the main prey of most mammalian mesopredators, and rodents are generally the alternative prey. We evaluated the bidirectional relation between the diel activity strategies of these mammalian mesopredators and prey coexisting in south‐western Europe. Results revealed that even though predation risk enforced by mammalian mesocarnivores during night‐time was approximately twice and five times higher than during twilight and daytime, respectively, murids consistently displayed unimodal nocturnal behaviour. Conversely, the European rabbits exhibited a bimodal pattern that peaked around sunrise and sunset. Despite the existence of some overlap between the diel rhythms of mesocarnivores and rabbits, their patterns were not synchronized. We suggest that the environmental stressors in our study areas are not severe enough to override the endogenous regulation of the circadian cycle in murids. European rabbits, however, are able to suppress their biological tendency for nocturnality by selecting a predominantly crepuscular pattern. In spite of the higher energetic input, mesocarnivores do not completely track rabbits’ activity pattern. They rather track rodents’ activity. We propose that these systems have probably evolved towards a situation where some degree of activity during high‐risk periods benefits the overall prey population survival, while the accessibility to sufficient prey prevents predators to completely track them.     

Phase-Synchronization of Daily Motor Activities Can Reveal Differential Circadian Patterns

A patient diagnosed with seasonal affective disorder (SAD) carried out prospective ratings of depression weekly for nearly a decade. A winter peak of depression and benzodiazepine intake was documented. However, over the years, the depressive episodes shifted toward spring in an apparent free-running circannual rhythm (periodogram peaks at 53 and 55 weeks). This patient may have an underlying seasonal propensity to depression no longer precisely entrained to environmental cues. (Chronobiology International, 18(2), 309–313, 2001)     

Influence of Seasonality on Activity Patterns, Feeding Behavior, Ranging, and Grouping Patterns in Taï Chimpanzees

Lowland rain forest chimpanzees (Pan troglodytes verus) of the Taï National Park, Ivory Coast, responded to the minor dry season (July and August) of 1988 in a predictable manner by spending more time feeding, feeding more frequently on lower quality food items, reducing day range and party size, and spending more time solitarily and less time in mixed groups than during the rainy season. These behaviors are consistent with a response to scarce resources. My findings do not support Boesch's (1991, 1996) hypothesis of bisexually bonded chimpanzees. Females spent 45% of time alone and associated with males in mixed parties only 18% of their time. This major discrepancy in our results probably stems from differences in the time of year when our studies were conducted, the year in which my study was conducted (potentially scarcer resources than on average), and methodological differences: focal animal sampling of males and females equally. Although Boesch (1991, 1996) and Steiner (1996) have demonstrated that Taï parties are usually larger and more mixed, Taï chimpanzee social structure—party size and composition—during this study closely resembles that found at other study sites.