EMG activity in hyoid muscles during pig suckling

Infant suckling is a complex behavior that includes cycles of rhythmic sucking as well as intermittent swallows. This behavior has three cycle types: 1) suck cycles, when milk is obtained from the teat and moved posteriorly into the valleculae in the oropharynx; 2) suck-swallow cycles, which include both a rhythmic suck and a pharyngeal swallow, where milk is moved out of the valleculae, past the larynx, and into the esophagus; and 3) postswallow suck cycles, immediately following the suck-swallow cycles. Because muscles controlling these behaviors are active in all three types of cycles, we tested the hypothesis that different patterns of electromyographic (EMG) activity in the mylohyoid, hyoglossus, stylohyoid, and thyrohyoid muscles of the pig characterized each cycle type. Anterior mylohyoid EMG activity occurred regularly in every cycle and was used as a cycle marker. Thyrohyoid activity, indicating the pharyngeal swallow, was immediately preceded by increased stylohyoid and hyoglossus activity; it divided the suck-swallow cycle into two phases. Timed from the onset of the suck-swallow cycle, the first phase had a relatively fixed duration while the duration of the second phase, timed from the thyrohyoid, varied directly with cycle duration. In short-duration cycles, the second phase could have a zero duration so that thyrohyoid activity extended into the postswallow cycle. In such cycles, all swallowing activity that occurred after the thyrohyoid EMG and was associated with bolus passage through the pharynx fell into the postswallow cycle. These data suggest that while the activity of some muscles, innervated by trigeminal and cervical plexus nerves, may be time locked to the cycle onset in swallowing, the cycle period itself is not. The postswallow cycle consequently contains variable amounts of pharyngeal swallowing EMG activity. The results exemplify the complexity of the relationship between rhythmic sucking and the swallow.     

The coordination and interaction between respiration and deglutition in young pigs

The anatomical pathways for inspired air and ingested food cross in the pharynx of mammals, implying that breathing and swallowing must be separated either in space or in time. In this study we investigated the time relationship between swallowing and respiration in young pigs, as a model for suckling mammals. Despite the high morphological position of the larynx in young mammals, allowing liquid to pass in food channels lateral to the larynx, respiration and swallowing are not wholly independent events. Although, when suckling on a veterinary teat, the swallows occurred at various points in the respiratory cycle, there was always a period of apnea associated with the swallow. Finally, an increase in the viscosity of the milk altered this coordination, changing respiratory cycle length and also restricting the relative rate at which swallows occurred in some parts of the respiratory cycle. These results suggest that the subsequent changes in respiratory activity at weaning, associated with passage of a solid bolus over the larynx, is preceded by the ability of the animal to alter coordination between respiration and swallowing for a liquid bolus. Accepted: 29 September 1997     

Mechanism of intraoral transport in macaques

All mammals have the same divisions of cyclic movement of tongue and hyoid during mastication: a protraction or forward phase that begins at minimum gape, and a retraction or return phase. Nonanthropoid mammals transport food from the oral cavity to the oropharynx during the return phase; food on the dorsal surface of the tongue moves distally while the tongue is retracted. Macaques, however, transport food during the protraction phase of tongue/hyoid movement. Food is squeezed posteriorly by contact between the tongue surface and the palate anterior to the food. This mechanism of transport is occasionally seen in nonanthropoid mammals when they are transporting liquids from the oral cavity to the oropharynx. It has, however, not been seen when these mammals transport solid food. One morphological basis for this difference is the reduction in height of the rugae of the palate of macaques. In most mammals these rugae are pronounced ridges that are able to hold food in place during protraction as the tongue slides forward beneath the food. Anthropoids and other mammals differ in the way they store food prior to swallowing. When macaques transport food to the oropharynx, usually they swallow in the next cycle, but always in the next 2 or 3 cycles. Most mammals transport and store food in the oropharynx for several cycles before a swallow clears that region of food. This behavior is correlated with differences in morphology of the oropharynx; anthropoids have reduced valleculae, the area in which other mammals store food prior to swallowing.     

Kinematics of the pharyngeal jaws during feeding in Oreochromis niloticus (Pisces,Perciformes)

Cichlids possess a complex pharyngeal jaw apparatus, the osteological components of which are two upper pharyngeal jaws, articulating with the neurocranial base, and a single lower pharyngeal jaw. Quantitative cinera-diography revealed that pharyngeal food processing in Oreochromis niloticus involves transport, mastication, and swallowing, effected by cyclical pharyngeal jaw movements. Transport and swallowing occur by simultaneous retractions of both upper pharyngeal jaws. Food reduction (mastication) is effected by lower jaw elevation (compression) and protraction (shear) during upper jaw retraction. Each movement cycle contains a transport, reduction, and swallowing component, although their relative importance may vary within a feeding sequence. The upper and lower pharyngeal jaws show opposite anteroposterior movements during most of the cycle. Variations in the amplitudes and the durations of the different movement components reflect the consistency and the size of the food.     

Effect of sensory input from the tongue on jaw movement in normal feeding in the opossum

Opossums were presented with solid and liquid foods. The movements of the jaw and tongue were recorded cineradiographically together with recordings of the EMG activity in muscles opening the jaw and moving the base of the tongue (hyoid). The jaw opening in each cycle was in two stages--01 and 02; 01 had a constant amplitude irrespective of the food ingested. Ingestion of liquid (which involved continuous accumulation of a liquid bolus in the valleculae prior to swallowing) was associated with cycles of oral movement in which 02 was small; tongue retraction was associated with this opening. In contrast, solid and semisolid food ingestion was associated with large angles of jaw opening in 02 that also coincided with the tongue retraction. In this latter case a characteristic pattern of EMG activity, in which all the muscles moving the hyoid were simultaneously active, was added to the pattern seen in lapping; this additional activity had an EMG pattern that was consistent with a jaw opening reflex. The findings contrast with other reports that the jaw opening reflex is suppressed in mastication. Experimentally induced tongue contact with a variety of solid surfaces during lapping (an activity involving accumulation of a liquid bolus in the valleculae) induced neither increased jaw opening nor the additional EMG pattern. However, in situations when there was no bolus in the valleculae, additional jaw opening activity was elicited when the tongue contracted solids intra- or extra-orally. It is suggested that the ability of sensory input, from the anterior tongue, to elicit a jaw opening reflex and to change the type of jaw/tongue cycle was dependent upon the extent of bolus accumulation in the valleculae and therefore indirectly upon the consistency of the food.     

Development of Nonsocial Behaviour in the Asiatic Elephant

The elephant calf, a defended follower is completely dependent on adults till the age of 3 months. It begins to explore and attempts to feed at 3 to 6 months, and then becomes partially independent with some feeding on its own. The characteristics of behavior in adults are examined and the development of this pattern in the calf is traced by analyis of duration, transition and clustering of the behavioural elements. Essential activities like suckling, locomotion and lying down for rest appear soon after birth whereas elements of feeding, grooming and play appear only at a later stage. The calf takes the initiative in suckling and its termination, drinking directly by mouth till the age of 6 months. The first element of feeding appears at about a week in attempts to pick up and hold objects in the trunk. Co-ordination of limb, trunk and mouth movement is achieved by about 1 month. The calf is strong enough to pull out plants by 6 months when independent feeding begins. In about a year feeding, drinking and dusting patterns are well developed.     

Respiration during feeding on solid food: alterations in breathing during mastication, pharyngeal bolus aggregation, and swallowing.

During feeding, solid food is chewed and propelled to the oropharynx, where the bolus gradually aggregates while the larynx remains open and breathing continues. The aggregated bolus in the valleculae is exposed to respiratory airflow, yet aspiration is rare in healthy individuals. The mechanism for preventing aspiration during bolus aggregation is unclear. One possibility is that alterations in the pattern of respiration during feeding could help prevent inhalation of food from the pharynx. We hypothesized that respiration was inhibited during bolus aggregation in the valleculae. Videofluorography was performed on 10 healthy volunteers eating solid foods with barium. Respiration was monitored concurrently with plethysmography and nasal air pressure. The timing of events during mastication, food transport, pharyngeal bolus aggregation, and swallowing were measured in relation to respiration. Respiratory cycle duration decreased during chewing (P < 0.001) but increased with swallowing (P < 0.001). During 66 recordings of vallecular bolus aggregation, there was inspiration in 8%, expiration in 41%, a pause in breathing in 17%, and multiple phases (including inspiration) in 35%. Out of 98 swallows, 47% started in the expiratory phase and 50% started during a pause in breathing, irrespective of bolus aggregation in the valleculae. Plethysmography was better than nasal manometry for determining the end of active expiration during feeding and swallowing with solid food. The hypothesis is rejected in that respiration was not inhibited during bolus aggregation. These findings suggest that airflow through the pharynx does not have a role in preventing aspiration during bolus aggregation in the oropharynx.     

The interaction between breathing and swallowing in healthy individuals

In this article, we aimed at investigating the interaction between breathing and swallowing patterns in normal subjects. Ten healthy volunteers were included in the study. Diaphragm EMG activity was recorded by a needle electrode inserted into the 7th or 8th intercostal space. Swallowing was monitored by submental EMG activity, and laryngeal vertical movement was recorded by using a movement sensor. A single voluntary swallow was initiated during either the inspiration or expiration phases of respiration, and changes in EMG activity were evaluated. When a swallow coincided with either inspiration or expiration, the duration of the respiratory phase was prolonged. Normal subjects were able to voluntarily swallow during inspiration. During the inspiration phase with swallowing, diaphragmatic activity did not ceased and during the expiration phase with swallowing, there was a muscle activity in the diaphragm muscle.     

Diurnal rhythms of hepatic carbohydrate metabolism during development of the rat

The ;8+16' feeding schedule (8h feeding and 16h without food in each 24h cycle) was applied to nursing mother rats to study enzyme development in neonatal rats in the absence of solid food. A ;16+8' suckling schedule (16h with the mother and 8h while the mother is fed in a separate cage) was used to show that the increases in pyruvate kinase, glucokinase and aldolase B activities that occur in the late suckling period of liver development do not require the intake of solid food at this time. Their activities may, however, be modulated by the composition of the diet at the time of weaning. Adaptation to the composition of the diet can occur within one feeding period, and to the periodicity of food provision in one or two feeding periods. In the early neonatal period, diurnal rhythms of tyrosine aminotransferase, liver glycogen and glucokinase are either greatly suppressed or absent, but develop rapidly after weaning. Food-dependent rhythms of glycogen and tyrosine aminotransferase were included in the late suckling period (day 14).     

The effect of age at metamorphosis on the transition from larval to adult suspension‐feeding of the slipper limpet Crepidula fornicata

Slipper limpets use different ciliary feeding mechanisms as larvae and adults. Veliger larvae of Crepidula fornicata developed part of the adult feeding apparatus, including ctenidial filaments, neck lobe, and radula, before metamorphosis, but ctenidial feeding did not begin until well after loss of the larval feeding apparatus (velum) at metamorphosis. Earlier initiation of ctenidial feeding by individuals that were older larvae when metamorphosis occurred suggests continued development toward ctenidial feeding during delay of metamorphosis. Early juveniles produced a ciliary current through the mantle cavity and moved the radula in a grasping action before they began to capture algal cells on mucous strands or form a food cord. Either early juveniles could not yet form mucous strands or they delayed their production until development of other necessary structures. The neck canal for transporting food from ctenidium to mouth cannot develop before velar loss. In their first feeding, juveniles fed much like the adults except that the neck canal was less developed and the path of the food cord toward the mouth sometimes varied. As suspension feeders, calyptraeids lack the elaborations of foregut that complicate transition to juvenile feeding for many caenogastropods, but a path for the food cord must develop after velar loss. Why individuals can initiate ctenidial feeding sooner when they are older at metamorphosis is not yet known. The juveniles became sedentary soon after metamorphosis and were not observed to feed by scraping the substratum with the radula, in contrast to the first feeding by juveniles of another calyptraeid species, observed by Montiel et al. ( 2005 ).     

Periodization of the early postnatal development in the rat with particular attention to the weaning period

The early postnatal period is characterized by a great plasticity with critical windows in which any inadequate insult or intervention may be able to influence both positively and adversely postnatal growth and development. After birth the rat littermates enter the presuckling period (initial 6 hours terminated by the first nursing), characterized by transition from the amniotic fluid to the air, by the changes of the ambient temperature, by the termination of placental nutrition and by oxidative stress. After this stage the suckling period initiates and the littermates start to consume milk of their mothers. Comsumption of milk culminates on day 15, then decreases and terminates on postnatal day 28. The end of the suckling period and the onset of physiological weaning is determined by the moment when the youngs for the first time consume the solid food together with milk (postnatal day 17 in rats). On day 19 the first intake of drinking water occurs. The weaning period terminates by the last consumption of maternal milk - on postnatal day 28. It is necessary to stress that the duration of early postnatal periods is independent of the changes of body weight. The precise knowledge of individual ontogenetic periods critical for further development is crucial for the prediction and explanation of reactions to various pathogenetic stimuli both under experimental conditions and in clinical medicine.     

Suppression of Abdominal Motor Activity during Swallowing in Cats and Humans

Diseases affecting pulmonary mechanics often result in changes to the coordination of swallow and breathing. We hypothesize that during times of increased intrathoracic pressure, swallow suppresses ongoing expiratory drive to ensure bolus transport through the esophagus. To this end, we sought to determine the effects of swallow on abdominal electromyographic (EMG) activity during expiratory threshold loading in anesthetized cats and in awake-healthy adult humans. Expiratory threshold loads were applied to recruit abdominal motor activity during breathing, and swallow was triggered by infusion of water into the mouth. In both anesthetized cats and humans, expiratory cycles which contained swallows had a significant reduction in abdominal EMG activity, and a greater percentage of swallows were produced during inspiration and/or respiratory phase transitions. These results suggest that: a) spinal expiratory motor pathways play an important role in the execution of swallow, and b) a more complex mechanical relationship exists between breathing and swallow than has previously been envisioned.     

Competition in size-structured populations: mechanisms inducing cohort formation and population cycles

In this paper we investigate the consequences of size-dependent competition among the individuals of a consumer population by analyzing the dynamic properties of a physiologically structured population model. Only 2 size-classes of individuals are distinguished: juveniles and adults. Juveniles and adults both feed on one and the same resource and hence interact by means of exploitative competition. Juvenile individuals allocate all assimilated energy into development and mature on reaching a fixed developmental threshold. The combination of this fixed threshold and the resource-dependent developmental rate, implies that the juvenile delay between birth and the onset of reproduction may vary in time. Adult individuals allocate all assimilated energy to reproduction. Mortality of both juveniles and adults is assumed to be inversely proportional to the amount of energy assimilated. In this setting we study how the dynamics of the population are influenced by the relative foraging capabilities of juveniles and adults.In line with results that we previously obtained in size-structured consumer-resource models with pulsed reproduction, population cycles primarily occur when either juveniles or adults have a distinct competitive advantage. When adults have a larger per capita feeding rate and are hence competitively superior to juveniles, population oscillations occur that are primarily induced by the fact that the duration of the juvenile period changes with changing food conditions. These cycles do not occur when the juvenile delay is a fixed parameter. When juveniles are competitively superior, two different types of population fluctuations can occur: (1) rapid, low-amplitude fluctuations having a period of half the juvenile delay and (2) slow, large-amplitude fluctuations characterized by a period, which is roughly equal to the juvenile delay. The analysis of simplified versions of the structured model indicates that these two types of oscillations also occur if mortality and/or development is independent of food density, i.e. in a situation with a constant juvenile developmental delay and a constant, food-independent background mortality. Thus, the oscillations that occur when juveniles are more competitive are induced by the juvenile delay per se. When juveniles exert a larger foraging pressure on the shared resource, maturation implies an increase not only in adult density, but also in food density and consequently fecundity. Our analysis suggests that this correlation in time between adult density and fecundity is crucial for the occurrence of population cycles when juveniles are competitively superior.     

Breast-feeding patterns, maternal milk output and lactational infecundity.

Breast feeding can serve as a contraceptive up to 6 months after birth with 98% effectiveness. The so-called suckling reflex arc is a sensitive mechanism conveying messages about nutritional needs of the child, stimulating the release of prolactin responsible for milk production, suppressing ovarian activity by hypothalamic beta-endorphin production leading to reduction in the pulsatile release of gonadotrophic hormones. Increased suckling produces more milk, but only in well-fed women. In Bangladesh, malnourished children required more suckling time than better-fed infants. Increasing the amounts of weaning food lowers feed urgency and suckling intensity thereby relaxing ovarian suppression. Breast-milk consumption can be measured by test weighing the infant before and after feeding, but it is subject to error, especially in developing countries where up to 20 feeds are required to deliver 800 ml of milk. Thus, weighing over 2-5 days is more accurate. The method of giving deuterium oxide tracer to the mother, and measuring its disappearance from the mother's milk and the infant's saliva for 2 weeks is precise and noninvasive. The Doppler ultrasound technique measures milk flow through an artificial teat for research purposes. Manual or mechanical pump measurement of milk transfer separates the mother from the child, thus it is not useful for normal conditions. It can be used for estimating the amount of milk left and the rate of milk secretion by using a breast pump along with oxytocin for full emptying of milk. Accurate estimation of suckling intensity and milk transfer is essential for the determination of the relationship of breast feeding and lactational infecundity.     

A comparison of food capture and ingestion in juveniles of two flatfish species, Pleuronectes platessa and Limanda limanda (Teleostei: Pleuronectiformes)

Juvenile plaice, Pleuronectes platessa ( n =6, 151–197 mm SL) and dab Limanda limanda ( n =6, 119–180.5 mm SL) feeding on pieces of mussel flesh were studied by high-speed (200 fields s⁻¹) videophotography at 11) C. Prey capture cycles (involving jaw protrusion and opercular expansion) were described in both species. The capture cycle was followed by at least one cycle (similar to the capture cycle in all respects except food intake) which is believed to transport food to the oesophagus. The transport cycle (or cycles) was followed by a species-specific number of normal respiratory cycles (7 or 8 in plaice, 11 or 12 in dab) and a final single cycle when food debris was violently ejected from the opercular cavity. The respiratory cycles and ejection cycle cleansed the orobranchial/opercular cavities and gills. Differences between the species were relatively minor, but plaice captured and transported food more quickly than dab.     

Linking stages of life history: How larval quality translates into juvenile performance for an intertidal barnacle (Balanus glandula)

Many marine invertebrates with complex life cycles produce planktoniclarvae that experience environmental conditions different fromthose encountered by adults. Factors such as temperature andfood, known to impact the larval period, can also affect larvalsize and consequently the size of newly settled juveniles. Afterdocumenting natural variation in the size of cyprids (the finallarval stage) of the barnacle Balanus glandula, we experimentallymanipulated temperature and food given to larvae to producecyprids of differing sizes but within the size range of cypridsfound in the field. In a set of trials in which larvae of B.glandula were raised on full or reduced rations in the laboratoryand subsequently outplanted into the field as newly metamorphosedjuveniles, we explored the effects of larval nutrition and sizeon juvenile performance. Larvae that received full rations throughouttheir feeding period produced larger cyprids (with more lipidand protein). These larger cyprids grew faster as juvenilesand sometimes survived better in the field than juveniles fromlarvae that had their food ration reduced in the last feedinginstar. For naturally settling barnacles brought into the laboratorywithin 2 days of settlement and fed, we found that initial juvenilesize was a good predictor of juvenile size even after 2 weeksof growth. By manipulating food given to juveniles that werederived from larvae fed either full or reduced rations, we foundthat larval nutritional effects persisted in juveniles for 2–3times the period that larvae experienced altered food rations.     

The use of the tongue and hyoid apparatus during feeding in lizards (Ctenosaura similis and Tupinambis nigropunctatus)

The use of the tongue and hyoid is examined in cineradiographic and electromyographic investigations of feeding in two species of lizards, Ctenosaura similis (Iguanidae) and Tupinambis nigropunctatus (Teiidae). In both animals food is transported through the oral cavity by regular cycles of the tongue. Tongue movements correlate with jaw and hyoid movement. Similarities between the two animals in the use of the tongue in food transport, lapping, pharyngeal packing, and pharyngeal emptying are detailed. Mechanisms of tongue protrusion are examined and it is shown that the tongue in Tupinambis is relatively more protrusible than in Ctenosaura. This difference is complementary with data on the greater reliance of Tupinambis on the tongue as a sensory organ. Tupinambis further differs from Ctenosaura in possessing a greater mobility of the hyoid. In many features of tongue use in food transport, lizards resemble mammals, supporting postulations of a basic pattern of intra-oral food transport. However, whether this pattern can be attributed to convergence or a common, primitive neural pattern of control cannot be distinguished. Lizards lack two major characteristics of mammalian food transport: regular masticatory cycles and an internal swallowing mechanism.     

Patterns of variation across primates in jaw-muscle electromyography during mastication

Biologists that study mammals continue to discuss the evolutionof and functional variation in jaw-muscle activity during chewing.A major barrier to addressing these issues is collecting sufficientin vivo data to adequately capture neuromuscular variation ina clade. We combine data on jaw-muscle electromyography (EMG)collected during mastication from 14 species of primates andone of treeshrews to assess patterns of neuromuscular variationin primates. All data were collected and analyzed using thesame methods. We examine the variance components for EMG parametersusing a nested ANOVA design across successive hierarchical factorsfrom chewing cycle through species for eight locations in themasseter and temporalis muscles. Variation in jaw-muscle EMGswas not distributed equally across hierarchical levels. Thetiming of peak EMG activity showed the largest variance componentsamong chewing cycles. Relative levels of recruitment of jawmuscles showed the largest variance components among chewingsequences and cycles. We attribute variation among chewing cyclesto (1) changes in food properties throughout the chewing sequence,(2) variation in bite location, and (3) the multiple ways jawmuscles can produce submaximal bite forces. We hypothesize thatvariation among chewing sequences is primarily related to variationin properties of food. The significant proportion of variationin EMGs potentially linked to food properties suggests thatexperimental biologists must pay close attention to foods givento research subjects in laboratory-based studies of feeding.The jaw muscles exhibit markedly different variance componentsamong species suggesting that primate jaw muscles have evolvedas distinct functional units. The balancing-side deep masseter(BDM) exhibits the most variation among species. This observationsupports previous hypotheses linking variation in the timingand activation of the BDM to symphyseal fusion in anthropoidprimates and in strepsirrhines with robust symphyses. The working-sideanterior temporalis shows a contrasting pattern with littlevariation in timing and relative activation across primates.The consistent recruitment of this muscle suggests that primateshave maintained their ability to produce vertical jaw movementsand force in contrast to the evolutionary changes in transverseocclusal forces driven by the varying patterns of activationin the BDM.