A biomechanical model for analysis of muscle force,power output and lower jaw motion in fishes

Fish skulls are complex kinetic systems with movable components that are powered by muscles. Cranial muscles for jaw closing pull the mandible around a point of rotation at the jaw joint using a third-order lever mechanism. The present study develops a lever model for the jaw of fishes that uses muscle design and the Hill equation for nonlinear length-tension properties of muscle to calculate dynamic power output. The model uses morphometric data on skeletal dimensions and muscle proportions in order to predict behavior and force transmission mediated by lever action. The computer model calculates a range of dynamic parameters of jaw function including muscle force, torque, effective mechanical advantage, jaw velocity, bite duration, bite force, work and power. A complete list of required morphometrics is presented and a software program (MandibLever 2.0) is available for implementing lever analysis. Results show that simulations yield kinematics and timing profiles similar to actual fish feeding events. Simulation of muscle properties shows that mandibles reach their peak velocity near the start of jaw closing, peak force at the end of jaw closing, and peak power output at about 25% of the closing cycle time. Adductor jaw muscles with different mechanical designs must have different contractile properties and/or different muscle activity patterns to coordinate jaw closing. The effective mechanical advantage calculated by the model is considerably lower than the mechanical advantage estimated from morphological lever ratios, suggesting that previous studies of morphological lever ratios have overestimated force and underestimated velocity transmission to the mandible. A biomechanical model of jaw closing can be used to interpret the mechanics of a wide range of jaw mechanisms and will enable studies of the functional results of developmental and evolutionary changes in skull morphology and physiology.     

Patterned jaw movements and the motor neuron activity during rejection of seaweed in Aplysia kurodai

Japanese Aplysia kurodai feeds well on Ulva. In the present experiments we collected several species of seaweed at a location with many animals and initially explored the preference behavior for them. The animals rejected Grateloupia, Pachydictyon, Gelidium and Laurencia with rhythmic jaw and radula movements (active rejection). The animals sometimes bit off a piece of them (biting-off response). Recording activity of muscles contributing to jaw-opening and jaw-closing in freely moving animals showed that the onset of the jaw-closing activity, which always started later than the jaw-opening activity during each cycle of ingestion of Ulva, was advanced toward that of the jaw-opening activity during each cycle of the active rejection. Semi-intact experiments also showed that application of Pachydictyon or Gelidium extract to the lip region advanced the firing onset of the jaw-closing motor neurons at the radula-retraction phase. Video analysis showed that during the Ulva response the jaws opened at the radula-protraction phase and remained half-open at the earlier radula-retraction phase, while the jaws opened similarly at the radula-protraction phase but immediately closed at the radula-retraction phase during the Pachydictyon or Gelidium response. Accepted: 8 October 1997     

Loading patterns and jaw movements during mastication in Macaca fascicularis: a bone-strain, electromyographic, and cineradiographic analysis

Rosette strain gage, electromyography (EMG), and cineradiographic techniques were used to analyze loading patterns and jaw movements during mastication in Macaca fascicularis. The cineradiographic data indicate that macaques generally swallow frequently throughout a chewing sequence, and these swallows are intercalated into a chewing cycle towards the end of a power stroke. The bone strain and jaw movement data indicate that during vigorous mastication the transition between fast close and the power stroke is correlated with a sharp increase in masticatory force, and they also show that in most instances the jaws of macaques are maximally loaded prior to maximum intercuspation, i.e. during phase I (buccal phase) occlusal movements. Moreover, these data indicate that loads during phase II (lingual phase) occlusal movements are ordinarily relatively small. The bone strain data also suggest that the duration of unloading of the jaw during the power stroke of mastication is largely a function of the relaxation time of the jaw adductors. This interpretation is based on the finding that the duration from 100% peak strain to 50% peak strain during unloading closely approximates the half-relaxation time of whole adductor jaw muscles of macaques. The EMG data of the masseter and medial pterygoid muscles have important implications for understanding both the biomechanics of the power stroke and the external forces responsible for the "wishboning" effect that takes place along the mandibular symphysis and corpus during the power stroke of mastication. Although both medial pterygoid muscles reach maximum EMG activity during the power stroke, the activity of the working-side medial pterygoid peaks after the balancing-side medial pterygoid. Associated with the simultaneous increase of force of the working-side medial pterygoid and the decrease of force of the balancing-side medial pterygoid is the persistently high level of EMG activity of the balancing-side deep masseter (posterior portion). This pattern is of considerable significance because the direction of force of both the working-side medial pterygoid and the balancing-side deep masseter are well aligned to aid in driving the working-side lower molars across the upper molars in the medial direction during unilateral mastication.(ABSTRACT TRUNCATED AT 400 WORDS)     

Jaw and hyolingual movements during prey transport in varanid lizards: effects of prey type

The ability to modulate feeding kinematics in response to prey items with different functional properties is likely a prerequisite for most organisms that feed on a variety of food items. Variation in prey properties is expected to reveal variation in feeding function and the functional role of the different phases in a transport cycle. Here we describe the kinematics of prey transport of two varanid species, Varanus niloticus and Varanus ornatus. These species were selected for analysis because of their highly specialised hyolingual system and food transport mechanism (inertial food transport). In these animals, tongue and hyoid movements are expected to make no, or only a minor, contribution to prey transport. We observed statistically significant prey type effects that could be associated with prey properties such as mass, size and mobility. These data show that both species are capable of modulating the kinematics of food transport in response to different prey types. Moreover, not only the kinematics of the jaws were modulated in response to prey characteristics but also the anterior/posterior movements of the tongue and hyoid. This suggests a more important role of the tongue and hyolingual movements in these animals than previously suspected. In contrast, head movements were rather stereotyped and were not modulated in response to changes in prey type.     

Functional morphology of bite mechanics in the great barracuda (Sphyraena barracuda)

The great barracuda, Sphyraena barracuda, is a voracious marine predator that captures fish with a swift ram feeding strike. While aspects of its ram feeding kinematics have been examined, an unexamined aspect of their feeding strategy is the bite mechanism used to process prey. Barracuda can attack fish larger than the gape of their jaws, and in order to swallow large prey, can sever their prey into pieces with powerful jaws replete with sharp cutting teeth. Our study examines the functional morphology and biomechanics of 'ram-biting' behavior in great barracuda where the posterior portions of the oral jaws are used to slice through prey. Using fresh fish and preserved museum specimens, we examined the jaw mechanism of an ontogenetic series of barracuda ranging from 20 g to 8.2 kg. Jaw functional morphology was described from dissections of fresh specimens and bite mechanics were determined from jaw morphometrics using the software MandibLever (v3.2). High-speed video of barracuda biting (1500 framess(-1)) revealed that prey are impacted at the corner of the mouth during capture in an orthogonal position where rapid repeated bites and short lateral headshakes result in cutting the prey in two. Predicted dynamic force output of the lower jaw nearly doubles from the tip to the corner of the mouth reaching as high as 58 N in large individuals. A robust palatine bone embedded with large dagger-like teeth opposes the mandible at the rear of the jaws providing for a scissor-like bite capable of shearing through the flesh and bone of its prey.     

Prey capture by Luciocephalus pulcher: implications for models of jaw protrusion in teleost fishes

Synopsis Luciocephalus pulcher possesses one of the most protrusible jaws known among teleosts, the premaxillae extending anteriorly a distance of 33% of the head length during feeding. Jaw bone movement during feeding proceeds according to a stereotypical pattern and resembles that of other teleosts except for extreme cranial elevation and premaxillary protrusion. Anatomical specializations associated with cranial elevation include: a highly modified first vertebra with a separate neural spine, articular fossae on the posterior aspect, greatly enlarged zygapophyses on the second vertebra with complex articular condyles, and highly pinnate multi-layered epaxial musculature with multiple tendinous insertions on the skull.Luciocephalus, despite the extreme jaw protrusion, does not use suction during prey capture: rather, the prey is captured by a rapid lunge (peak velocity of about 150 cm per sec) and is surrounded by the open mouth. Previous hypotheses of the function of upper jaw protrusion are reviewed in relation to jaw movements inLuciocephalus. Protrusion is not obligatorily linked with suction feeding; behavioral aspects of the feeding process limit the possible range of biological roles of a given morphological specialization, and make prediction of role from structure risky.     

Factors underlying the perturbation resistance of the trunk in the first part of a lifting movement

In the first part of lifting movements, the trunk movement is surprisingly resistant to perturbations. This study examined which factors contribute to this perturbation resistance of the trunk during lifting. Three possible mechanisms were studied: force-length-velocity characteristics of muscles, the momentum of the trunk as well as the effect of passive extending of the elbows. A forward dynamics modelling and simulation approach was adopted with two different input signals: (1) stimulation of Hill-type muscles versus (2) net joint moments. Experimental data collected during an unperturbed lifting movement were used as a reference, which a simulated lifting movement had to resemble. Subsequently, the simulated lifting movement was perturbed by applying 10 kg extra mass at the wrist (both before and after lift-off and with/without a fixed elbow), without modifying the input signals. The momentum of the trunk appeared to be insufficient to explain the perturbation resistance of trunk movements as found experimentally. In addition to the momentum of the trunk, the force-length-velocity characteristics of the muscles are necessary to account for the observed perturbation resistance. Initial extension of the elbow due to the mass perturbation delayed the propagation of the load to the shoulder. However, this delay is reduced due to the impedance at the elbow provided by the characteristics of muscles spanning the elbow. So, the force-length-velocity characteristics of the muscles spanning the elbow joint increase the perturbation at the trunk.     

The roles of head movements in the search and capture strategy of a tern (Aves, Laridae)

 Gull-billed terns (Gelochelidon nilotica) were video-filmed while searching for and capturing fiddler crabs. Search consists of a vertical head nystagmus, with fast upward flicks and downward slow phases made at the angular speed of the substrate in the approximate direction of the bill. The bill points down at about 60° during hunting, but is brought up to 15° from time to time, which brings the visual streak into line with the horizon; 45° roll movements of the head are consistent with alternation between the use of the temporal and central foveas to view the same object. When a crab has been detected the nystagmus is suspended, and the tern tracks the crab continuously as it manoeuvres into a catching position. This may involve tucking the head under the body so that the bill is 45° behind the vertical, and flying up and backwards for some metres, straightening up the head at the same time. Accepted: 7 November 1998     

Effects of bisphosphonates on keratinocytes and fibroblasts having a role in the development of osteonecrosis of the jaw

Bisphosphonates (BPs) have became the treatment of choice for the prevention of skeletal complications in cancer patients with bone metastases as well as in patients suffering from osteoporosis, Paget's disease and rheumatoid arthritis. Osteonecrosis of the jaw (ONJ) is a recently described complication associated with the use of BPs in which the key finding is exposed necrotic bone in the oral cavity. Often, the precipitating event appears to be a dental invasive procedure. We recently provided evidence that ONJ is associated with dental extractions and use of dentures. It has been reported that the primary lesion lies in the bone and it is related to over-suppression of bone turnover, but it is unclear why such a lesion should present with loss of the soft tissue covering the jawbone. We propose that BP could be impairing molecular signalling not only of osteoblasts and osteoclasts but also of fibroblasts and keratinocytes, via cell to cell endocrine and paracrine interactions in a double manner. Such an impairment would result to fibroblast and keratinocyte impaired multiplication, proliferation and migration thereby leading to defective mucosal wound healing. This provides an open entry point for the oral flora to reach the underlying jawbone which is considered to have poor metabolic and immune properties when under BP treatment. We demonstrate that ONJ is associated with mucosal damage, which could be mediated via BP induced soft tissue toxicity. BPs have been reported to promote keratinocyte and fibroblast apoptosis and to impair various cellular activities like apoptosis, RANK, RANK-L and OPG signalling, bone morphogenetic protein signalling, growth factor signalling, immune homeostasis and wound healing. We discuss potential consequences of the above hypothesis for practitioners and investigators.     

Fetal movements during late gestation in the pig: A longitudinal ultrasonographic study

Repeated ultrasonographic observation of fetal movements was used to distinguish movement patterns and to investigate the rate of occurrence and temporal organisation of these patterns (rest-activity cycles) during the last three weeks of gestation in the pig.By means of transabdominal ultrasonography with a 3.5 MHz linear array transducer, motility in ten different fetuses (one per sow) was studied. Six (median; range 4-6) 1 h recordings were made per fetus at 3-5 day intervals. Fifty-five 1 h recordings were available for analysis. The occurrence of fetal general movements (GM), isolated head (HM), forelimb movements (LM), and rotations (ROT) was analysed from video tapes. For each movement pattern, the trend in occurrence over time was assessed by multilevel analysis. The temporal association between different movement patterns was studied by calculation of the kappa value.ROT occurred very infrequently and showed no particular trend over time. GM, HM, and LM showed a significant decreasing trend towards parturition (P < 0.01). Total fetal activity (i.e., the sum of the four movement incidences) declined from an average of 25% of recording time to 9% over the last three weeks of pregnancy. Periods of fetal quiescence gradually increased with progressing gestation (P < 0.05). There was no evidence of concordant association between the periods of rest and activity of GM, HM, and LM or of improved temporal linkage between these movement patterns with time.Fetal bodily activity decreases towards parturition mainly due to prolonged periods of rest. Fetal movement patterns show rest-activity cycles, but each pattern appears to cycle independently from the other throughout late gestation. The present results of spontaneous fetal movements in the pig provide reference data for future studies of fetal activity under different zootechnical conditions or pharmacological interventions.     

A multisegmental cross-bridge kinetics model of the myofibril

Striated muscle is a mechanical system that develops force and generates power in serving vital activities in the body. Striated muscle is a complex biological system; a single mammalian muscle fibre contains up to hundred or even more myofibrils in parallel connected via an inter-myofibril filament network. In one single myofibril thousands of sarcomeres are lined up as a series of linear motors. We recently demonstrated that half-sarcomeres (hS) in a single myofibril operate non-uniformly. We outline a mathematical framework based on cross-bridge kinetics for the simulation of the force response and length change of individual hS in a myofibril. The model describes the muscle myofibril in contraction experiments under various conditions. The myofibril is modeled as a multisegmental mechanical system of hS models, which have active and viscoelastic properties. In the first approach, a two-state cross-bridge formalism relates the hS force to the chemical kinetics of ATP hydrolysis, as first described by Huxley [1957. Muscle structure and theories of contraction. Prog. Biophys. Mol. Biol. 7, 255-318]. Two possible types of biological variability are introduced and modeled. Numerical simulations of a myofibril composed of four to eight hS show a non-uniform hS length distribution and complex internal dynamics upon activation. We demonstrate that the steady-state approximation holds only in restricted time zones during activation. Simulations of myofibril contraction experiments that reproduce the classic steady-state force-length and force-velocity relationships, strictly constrained or “clamped” in either end-held isometric or isotonic contraction conditions, reveal a small but conspicuous effect of hS dynamics on force.     

The role of the frontal ganglion in foregut movements of the moth,Manduca sexta

Two types of rhythmic foregut movements are described in fifth instar larvae of the moth, Manduca sexta. These consist of posteriorly-directed waves of peristalsis which move food toward the midgut, and synchronous constrictions of the esophageal region, which appear to retain food within the crop. We describe these movements and the muscles of the foregut that generate them.The firing patterns of a subset of these muscles, including a constrictor and dilator pair from both the esophageal and buccal regions of the foregut, are described for both types of foregut movement.The motor patterns for the foregut muscles require innervation by the frontal ganglion (FG), which lies anterior to the brain and contains about 35 neurons. Eliminating the ventral nerve cord, leaving the brain and FG intact, did not affect the muscle firing patterns in most cases. Eliminating both the brain and the ventral nerve cord, leaving only the FG to innervate the foregut, generally resulted in an increased period for both gut movements and muscle bursts. This manipulation also produced increases in burst durations for most muscles, and had variable effects on the phasing of muscle activity. Despite these changes, the foregut muscles still maintained a rhythmic firing pattern when innervated by the FG alone.Two nerves exit the FG to innervate the foregut musculature: the anteriorly-projecting frontal nerve, and the posteriorly-directed recurrent nerve. Cutting the frontal nerve immediately and irreversibly stopped all muscle activity in the buccal region, while cutting the recurrent nerve immediately stopped all muscle activity in the pharyngeal and esophageal regions. Recordings from the cut nerves leaving the FG showed that the ganglion was spontaneously active, with rhythmic activity continuing within the nerves. These observations indicate that all of the foregut muscle motoneurons are located within the FG, and the FG in isolation produces a rhythmic firing pattern in the motoneurons. We have identified several motoneurons within the FG, by cobalt backfills and/or simultaneous intracellular recordings and fills from putative motoneurons and their muscles.Abbreviations BC Buccal Constrictor - BC1 buccal constrictor motoneuron 1 - BC2 buccal constrictor motoneuron 2 - BD Buccal Dilator - BD1 buccal dilator motoneuron 1 - EC Esophageal Dilator - EC1 esophageal dilator motoneuron 1 - EC2 esophageal dilator motoneuron 2 - EC3 esophageal dilator motoneuron 3 - ejp excitatory junction potential - FG frontal ganglion - psp postsynaptic potential     

The feeding ecology of the pit-making ant lion larva,Myrmeleon bore: Feeding rate and species composition of prey in a habitat

The prey species composition and feeding rate of the pit-making ant lion larva,Myrmeleon bore Tjeder, which inhabits open sandy areas, were examined. Not less than 30 prey species, most of which were ants, were collected during a research period of 1.5 years. First instar larvae most often (81.1%) captured ants. Although 3rd instar larvae captured larger-sized prey than individuals of any other instar, they also captured small prey. The feeding rate of 3rd instar larvae was estimated by using the frequency of observed predation (FOP; (no. of ant lions handling a prey)/(total no. of pits observed)), the prey-handling time and the rhythm of daily foraging activity. FOP ofM. bore larvae was constant on the whole from spring to autumn. It was estimated that each captured 1.25 prey per day on average during this period. This estimate, however, was the feeding rate for days on which there was no rain. Assuming that the larvae cannot capture prey due to pit destruction when there is more than 10 mm of rainfall per day, the figure was reduced to 1.03 prey/day. The estimated feeding rate was evaluated with reference to larval foraging behavior.     

A forceful upper jaw facilitates picking-based prey capture: biomechanics of feeding in a butterflyfish,Chaetodon trichrous

Biomechanical models of feeding mechanisms elucidate how animals capture food in the wild, which, in turn, expands our understanding of their fundamental trophic niche. However, little attention has been given to modeling the protrusible upper jaw apparatus that characterizes many teleost species. We expanded existing biomechanical models to include upper jaw forces using a generalist butterflyfish, Chaetodon trichrous (Chaetodontidae) that produces substantial upper jaw protrusion when feeding on midwater and benthic prey. Laboratory feeding trials for C. trichrous were recorded using high-speed digital imaging; from these sequences we quantified feeding performance parameters to use as inputs for the biomechanical model. According to the model outputs, the upper jaw makes a substantial contribution to the overall forces produced during mouth closing in C. trichrous. Thus, biomechanical models that only consider lower jaw closing forces will underestimate total bite force for this and likely other teleost species. We also quantified and subsequently modeled feeding events for C. trichrous consuming prey from the water column versus picking attached prey from the substrate to investigate whether there is a functional trade-off between prey capture modes. We found that individuals of C. trichrous alter their feeding behavior when consuming different prey types by changing the timing and magnitude of upper and lower jaw movements and that this behavioral modification will affect the forces produced by the jaws during prey capture by dynamically altering the lever mechanics of the jaws. In fact, the slower, lower magnitude movements produced during picking-based prey capture should produce a more forceful bite, which will facilitate feeding on benthic attached prey items, such as corals. Similarities between butterflyfishes and other teleost lineages that also employ picking-based prey capture suggest that a suite of key behavioral and morphological innovations enhances feeding success for benthic attached prey items.     

Body movements and their role as triggers of heartbeats in pupae of the Colorado potato beetle Leptinotarsa decemlineata

Rhythmic body movements and their role as triggers of intermittent heartbeats were studied in pupae of Leptinotarsa decemlineata Say. Heartbeats and body movements were recorded simultaneously by means of an optical method (infra‐red cardiography) combined with respirometry. IR‐cardiography allows heartbeats and body movements to be distinguished on the basis of their different rates (40–80 min^?1 and 4–8 min^?1, respectively) and amplitude. In the mid‐interecdysial period, abdominal movements in the pupae were always accompanied by heart activity beginning after the first 5–12 abdominal strokes. Simultaneous periods of abdominal movements and heartbeats lasted 2–5 min, while the intervening pause ranged from 40 to 72 min at 24°C. Experiments of forced heart activity showed that a slight external tactile stimulus (prodding once with a single hair), applied prior to an expected heartbeat bout, evoked abdominal movements followed soon afterwards by contractions of the dorsal vessel. Repeated prodding with a hair evoked body rotating movements (1–3 strokes) at any time with heartbeats starting at the first movement. We surmise that rhythmic body movements play an active role in the triggering of heart activity in pupae according to the principle of cardiac reflex response. This is a possible mechanism for synchronizing periods of heart activity with periods of rhythmic body movements. Haemolymph circulation in pupae is favoured when heartbeat bouts are accompanied by active body movements.     

A fuzzy-genetic model for estimating forces from electromyographical activity of antagonistic muscles due to planar lower arm movements: the effect of nonlinear muscle properties

The aim of this paper is to create a model for mapping the surface electromyogram (EMG) signals to the force that generated by human arm muscles. Because the parameters of each person's muscle are individual, the model of the muscle must have two characteristics: (1) The model must be adjustable for each subject. (2) The relationship between the input and output of model must be affected by the force-length and the force-velocity behaviors are proven through Hill's experiments. Hill's model is a kinematic mechanistic model with three elements, i.e. one contractile component and two nonlinear spring elements.In this research, fuzzy systems are applied to improve the muscle model. The advantages of using fuzzy system are as follows: they are robust to noise, they prove an adjustable nonlinear mapping, and are able to model the uncertainties of the muscle.Three fuzzy coefficients have been added to the relationships of force-length (active and passive) and force-velocity existing in Hill's model. Then, a genetic algorithm (GA) has been used as a biological search method that can adjust the parameters of the model in order to achieve the optimal possible fit.Finally, the accuracy of the fuzzy genetic implementation Hill-based muscle model (FGIHM) is invested as following: the FGIHM results have 12.4% RMS error (in worse case) in comparison to the experimental data recorded from three healthy male subjects. Moreover, the FGIHM active force-length relationship which is the key characteristics of muscles has been compared to virtual muscle (VM) and Zajac muscle model. The sensitivity of the FGIHM has been evaluated by adding a white noise with zero mean to the input and FGIHM has proved to have lower sensitivity to input noise than the traditional Hill's muscle model.     

The stabilizing role of the Sabbath in pre-monarchic Israel:a mathematical model

The three monotheistic cultures have many common institutions and some of them germinated in pre-monarchic Israel. Reasonably, the essential institutions were in place at that starting point; this work explores the possibility that the Sabbath is one of these institutions. Our mathematical examination points to the potential cultural, civic, and social role of the weekly Sabbath, that is, the Sabbath institution, in controlling deviation from social norms. It begins with an analogy between spread of transgression (defined as lack of conformity with social norms) and of biological infection. Borrowing well-known mathematical methods, we derive solution sets of social equilibrium and study their social stability. The work shows how a weekly Sabbath could in theory enhance social resilience in comparison with a similar assembly with a more natural and longer period, say between New Moon and Full Moon. The examination reveals that an efficient Sabbath institution has the potential to ensure a stable organization and suppress occasional appearances of transgression from cultural norms and boundaries. The work suggests the existence of a sharp threshold governed by the “Basic Sabbath Number ש₀”—a critical observance of the Sabbath, or large enough ש₀, is required to ensure suppression of transgression. Subsequently, the model is used to explore an interesting question: how old is the Sabbath? The work is interdisciplinary, combining anthropological concepts with mathematical analysis and with archaeological parallels in regards to the findings.

Electronic supplementary material

The online version of this article (doi:10.1007/s10867-014-9373-9) contains supplementary material, which is available to authorized users.     

A dynamic model of water gain and loss in larval Mexican bean beetle

Abstract. Quantitative effects of temperature, vapour pressure deficit, host, and larval body size were experimentally determined. A simulation model for dynamic water balance in the Mexican bean beetle, Epilachna varivestis Mulsant, is presented and parameters are estimated from laboratory data for water gain/loss equations. The model is based on water loss through the cuticle, spiracles and frass, and water gain through ingestion.     

Tooth movements are guided by specific contact areas between the tooth root and the jaw bone: A dynamic 3D microCT study of the rat molar

Teeth sustain high loads over a lifetime and yet intact tooth failure is rare. The different structures of the tooth, jaw bone and the intervening soft periodontal ligament enable the tooth to endure repeated loading during mastication. Although mechanical and functional properties of the different components are thoroughly investigated, the manner in which the whole tooth functions under load is still enigmatic. A custom-made loading system inside a microCT scanner was used to directly visualize the root movements in relation to the jaw bone as the rat molar tooth was loaded. At low loads no contact was observed between the root surface and the bone, whereas at higher loads three specific contact areas between the root surface and the jaw bone were observed. These contact areas restrict tooth movement in the buccal-lingual direction, but enable the tooth to rock in a "seesaw" like manner in the distal-mesial direction. The contact areas appear to play a role in determining tooth motion and in turn define the manner in which the whole tooth moves when loaded. These observations are important for understanding basic structure-function relations of the tooth-PDL-bone system, and have direct implications for better understanding pathological and therapeutic processes in orthodontics, periodontics and jaw bone regeneration.