Mastication-related neurons in the orofacial first somatosensory cortex of awake cats

In the orofacial area of the first somatosensory cortex (SI), we recorded single unit activity from 699 neurons in 11 awake cats. Fifty-two percent (362/699) were mastication-related neurons (MRNs) showing activity related to some aspects of masticatory movements. MRNs were divided into three types by their activity patterns: (1) the rhythmical type, showing rhythmical bursts in pace with the masticatory rhythm; (2) the sustained type, showing a sustained firing during the period of taking food and (3) the transient (biting) type, showing intense discharges in coincidence with biting hard food. MRNs had mechanoreceptive fields in the perioral, tongue, periodontal and mandibular regions. The activities of perioral rhythmical-MRNs, mandibular transient-MRNs, tongue rhythmical-MRNs and periodontal transient-MRNs were correlated with food texture, while perioral rhythmical-MRNs, perioral sustained-MRNs and tongue sustained-MRMs were not. Both facial and intraoral MRNs were scattered throughout the facial and intraoral projection areas in SI. These findings provide evidence that the orofacial SI monitors masticatory movements for food ingestion.     

Classification of lumbosacral neurons by their discharge pattern during evoked locomotion

Unit activity was recorded in the lumbosacral division of the spinal cord during evoked locomotion in mesencephalic cats with the afferent fibers from their hind limbs intact or divided. If the afferent fibers were intact, all neurons recorded showed modulation of activity during locomotion in the rhythm of stepping movements. In experiments on cats with afferent fibers from the hind limbs divided modulation was absent in 30% of neurons, while in the modulated neurons, the frequencies in the excitation phase were approximately the same as when the limb innervation was intact. Modulation of activity in some neurons occurred in response to stimulation of the locomotor region even before stepping movements began. The tuning of the spinal generator of stepping movements is discussed.M. V. Lomonosov Moscow State University. Institute of Problems in Information Transmission, Academy of Sciences of the USSR, Moscow. Translated from Neirofiziologiya, Vol. 4, No. 4, pp. 410–417, July–August, 1972.     

Corticospinal mirror neurons

Here, we report the properties of neurons with mirror-like characteristics that were identified as pyramidal tract neurons (PTNs) and recorded in the ventral premotor cortex (area F5) and primary motor cortex (M1) of three macaque monkeys. We analysed the neurons’ discharge while the monkeys performed active grasp of either food or an object, and also while they observed an experimenter carrying out a similar range of grasps. A considerable proportion of tested PTNs showed clear mirror-like properties (52% F5 and 58% M1). Some PTNs exhibited ‘classical’ mirror neuron properties, increasing activity for both execution and observation, while others decreased their discharge during observation (‘suppression mirror-neurons’). These experiments not only demonstrate the existence of PTNs as mirror neurons in M1, but also reveal some interesting differences between M1 and F5 mirror PTNs. Although observation-related changes in the discharge of PTNs must reach the spinal cord and will include some direct projections to motoneurons supplying grasping muscles, there was no EMG activity in these muscles during action observation. We suggest that the mirror neuron system is involved in the withholding of unwanted movement during action observation. Mirror neurons are differentially recruited in the behaviour that switches rapidly between making your own movements and observing those of others.     

Rhythmically firing (20–50 Hz) neurons in monkey primary somatosensory cortex: Activity patterns during initiation of vibratory-cued hand movements

The activity patterns of rhythmically firing neurons in monkey primary somatosensory cortex (SI) were studied during trained wrist movements that were performed in response to palmar vibration. Of 1,222 neurons extracellularly recorded in SI, 129 cells (11%) discharged rhythmically (at 30 Hz) during maintained wrist position. During the initiation of vibratory-cued movements, neuronal activity usually decreased at 25 ms after vibration onset followed by an additional decrease in activity at 60 ms prior to movement onset. Rhythmically firing neurons are not likely to be integrate-and-fire neurons because, during activity changes, their rhythmic firing pattern was disrupted rather than modulated. The activity pattern of rhythmically firing neurons was complimentary to that of quickly adapting SI neurons recorded during the performance of this task (Nelson et al., 1991). Moreover, disruptions of rhythmic activity of individual SI neurons were similar to those reported previously for local field potential (LFP) oscillations in sensorimotor cortex during trained movements (Sanes and Donoghue, 1993). However, rhythmic activity of SI neurons did not wax and wane like LFP oscillations (Murthy and Fetz, 1992; Sanes and Donoghue, 1993). It has been suggested that fast (20–50 Hz) cortical oscillations may be initiated by inhibitory interneurons (Cowan and Wilson, 1994; Llinas et al., 1991; Stern and Wilson, 1994). We suggest that rhythmically firing neurons may tonically inhibit quickly adapting neurons and release them from the inhibition at go-cue onsets and prior to voluntary movements. It is possible that rhythmically active neurons may evoke intermittent oscillations in other cortical neurons and thus regulate cortical population oscillations.     

Neuronal activity of the monkey striatum points at the integration of successive actions into functional blocks

In order to study the role of the striatum in generation of multistage behavior, the spike activity of 148 cells was recorded in the monkey brain putamen. Two kinds of neuron responses were observed. Phasic response involved activity during only one stage of the behavior program, and tonic response involved activity during more than one sequential stage. The tonic responses were recorded in 132 neurons out of 148, 11 neurons responding only as tonic. Other 121 cells show under different conditions both tonic and phasic responses. Beginnings and ends of "tonic" responses as a rule corresponded to the start and completion of the nearest behavioral aim. The obtained data suggest that the neuron activity of striatum is related not only to the control of individual movements but also to the whole structure of behavior.     

Effects of food motivation on spike activity of cat somatic cortex neurons

Spike response to unconditioned electrocutaneous stimulation was investigated in cortical neurons of areas 3 and 4 in untrained hungry cats during heightened excitation motivated by food presentation and when at rest. This reinforcement led to changed background activity level, reduced intensity of the initial stages of spike response, and disappearance of late neuronal response. Neuronal response of the same cortical area to a conditioned stimulus (a clicking sound) during reduced food motivation (the animals being sated during the course of the experiment) was also studied under the effects of instrumental feeding reflex. Coordination between the timing of neuronal response and their corresponding movements was discovered from comparing response pattern accompanying the execution of paw-placing (conditioned reflex and intersignal) movements and those recorded at different levels of food-induced excitation, as well as a similarity between these reactions. It was found that the initial stages of neuronal response to a conditioned signal only occurred during contraction of the brachial biceps muscle, while coordination between their timing and that of EMG changes was dependent on the animal's degree of satiation. Findings indicate the possibility of food-induced excitation substantially influencing spike response pattern in somatic cortex neurons.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 19, No. 6, pp. 725–735, November–December, 1987.     

Patterns of spatio-temporal correlations in the neural activity of the cat motor cortex during trained forelimb movements

In order to study how neurons in the primary motor cortex (MI) are dynamically linked together during skilled movement, we recorded simultaneously from many cortical neurons in cats trained to perform a reaching and retrieval task using their forelimbs. Analysis of task-related spike activity in the MI of the hemisphere contralateral to the reaching forelimb (in identified forelimb or hindlimb representations) recorded through chronically implanted microwires, was followed by pairwise evaluation of temporally correlated activity in these neurons during task performance using shuffle corrected cross-correlograms. Over many months of recording, a variety of task-related modulations of neural activities were observed in individual efferent zones. Positively correlated activity (mainly narrow peaks at zero or short latencies) was seen during task performance frequently between neurons recorded within the forelimb representation of MI, rarely within the hindlimb area of MI, and never between forelimb and hindlimb areas. Correlated activity was frequently observed between neurons with different patterns of task-related activity or preferential activity during different task elements (reaching, feeding, etc.), and located in efferent zones with dissimilar representation as defined by intracortical microstimulation. The observed synchronization of action potentials among selected but functionally varied groups of MI neurons possibly reflects dynamic recruitment of network connections between efferent zones during skilled movement.     

Comparison of neural activity in the supplementary motor area and in the primary motor cortex in monkeys.

Neuronal activity recorded from the primary motor cortex (MI) and from the supplementary motor area (SMA) was compared in two monkeys trained to perform conditioned arm movements. A handle had to be held in a central waiting position until a visual go and cueing signal indicated to the monkey to move the handle either to a medial or to a lateral target zone (choice reaction time paradigm). Unit and representative electromyographic data were analyzed in relation either to the go signal or to movement onset. In 240 penetrations, 431 SMA neurons and 353 MI neurons were found with activity related to the task. The majority of neurons (303 in MI, 290 in SMA) displayed activity changes after the go signal and before movement onset. Of these "short-lead neurons", 71% in MI and 41% in SMA were clearly related to movement execution. The distribution of lead times in MI and SMA neurons was completely overlapping without any statistical difference among subgroups. The remaining neurons were as well related to the go signal as to movement onset, or were better related to the visual go signal. The response latencies to this signal were not statistically different in SMA and MI neurons. Activity changes during the waiting period was observed more frequently in SMA (47%) than in MI (32%); modulations restricted to the waiting period occurred in 14% of SMA neurons, but were exceptional in MI neurons (3%). It is concluded from these experiments that a surprisingly large proportion of SMA neurons have "MI-like" properties, in that they are temporally recruited together with MI neurons, with similar patterns of discharges during the task. This then suggests that the two interconnected areas operate in parallel. A population of SMA neurons is involved in some processing that is not as predominantly expressed in MI. This activity could relate to sensory, timing, or other higher-order aspects of response preparation, and/or motor functions such as postural stabilization.     

Activity of premotor vestibular neurons in the alert cat

The activity of medial vestibular nucleus neurons projecting to the contralateral abducens nucleus (premotor vestibular neurons) has been recorded during spontaneous and vestibular induced eye movements in the alert cat. Recorded neurons were identified by their antidromic activation from the abducens nucleus and by the post-synaptic field potential induced in this nucleus. The activity of identified medial vestibular neurons increased significantly with horizontal eye position and velocity toward the contralateral side, and decreased abruptly during ipsilateral saccades. The activity of these neurons was also related to head velocity toward the ipsilateral side. The functional role and origin of eye position and velocity signals present in these vestibular neurons are discussed.     

The oscillatory system responsible for the oculomotor activity during the bursts of REM.

1. In precollicular decerebrate cats the electrical activity of single pontine neurons was recorded before, during and after the episodes of postural atonia produced by i.v. injection of 0.03-0.1 mg/kg of eserine sulphate. These episodes were characterized by the regular occurrence of horizontal conjugate eye movements, which were mainly grouped in bursts of REM; moreover, a burst of REM in one direction was generally followed by a burst of REM in the opposite direction. 2. Among the recorded units, 32 showed an increase in their discharge rate during these cataplectic episodes. However, while these units fired at regular frequency when postural rigidity was present, they showed periodic changes in their discharge rate as soon as the bursts of REM appeared in the electrooculogram. In particular a nearly sinusoidal increase in the discharge rate was related to the appearance of an ocular burst in one direction, while a decrease in the unit discharge occurred during an ocular burst in the opposite direction. In some instances neighbouring pontine units located within each side of the brain stem showed reciprocal rate profiles during REM bursts oriented in a given direction, making it likely that the cyclic alternation of their activity depended upon their reciprocal interaction. 3. The alternative hypothesis, i.e., that these periodic changes in unit discharge depend upon the proprioceptive feedback due to the eye movements was excluded by the fact that these changes started before the occurrence of the bursts of REM and began to decline before the end of the burst. Moreover no variation in their firing rate was observed during the positional nystagmus induced by tilting the animal in the control period, i.e., when postural rigidity had reappeared following the end of the cataplectic episode. 4. Most of the neurons showing periodic changes in their discharge frequency during the bursts of REM were located in the pontine reticular formation. Scattered units were also found within the region of the locus coeruleus and the raphe system, close to the surrounding reticular structures. 5. In addition to these neurons, 60 pontine units were recorded, which did not show any changes in their discharge rate during transition from the control period to the cataplectic episode. However, phsiic increases or phasic decreases in their discharge rate appeared synchronously with the individual eye movements. Since in most instances these phasic changes in unit activity coincided with the appearance of the individual monophasic potentials recorded from the ascending MLB, which immediately preceded the rapid eye movements, these units could be attributed either to the premotor neurons responsible for these REM or to the closely related structures which generate their rhythmic discharge. In only a few instances did the discharge of these units not precede but follow the individual eye movements, indicating that they resulted from a proprioceptive feedback originating during the eye movements. 6...     

Response of recticular and dorsal thalamic nuclei neurons during extinction of conditioned implementation in the cat

Unit activity in 66 neurons of the reticular (R) nucleus and 31 neurons of the ventropostrolateral nuclei of the thalamus, and 14 neurons of the posterolateral nuclear complex, the pulvinar, were studied during extinction of the conditioned food implementation reflex. The number of R neurons that had responded to initial excitation in the first 300 msec after the conditional stimulus (CS) decreased with the extinction. Simultaneous disappearance of conditioned-reflex placement movements and late excitatory and inhibitory responses of R and dorsal thalamic nuclei neurons with latent periods exceeding 300 msec was also observed. Extinction of the conditioned reflex (CR) led to a significant lowering of background activity in two-thirds of investigated R and other thalamic nuclear neurons. This suggests that efferent effects from the reticular nucleus are decreased during Cr extinction.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the USSR, Kiev. Translated from Neirofiziologiya, Vol. 23, No. 1, pp. 3–8, January–February, 1991.     

Dispersed activity during passive movement in the globus pallidus of the 1-methyl-4-phenyl-1,2,3,6-tetrahydropyridine (MPTP)-treated primate

Parkinson's disease is a neurodegenerative disorder manifesting in debilitating motor symptoms. This disorder is characterized by abnormal activity throughout the cortico-basal ganglia loop at both the single neuron and network levels. Previous neurophysiological studies have suggested that the encoding of movement in the parkinsonian state involves correlated activity and synchronized firing patterns. In this study, we used multi-electrode recordings to directly explore the activity of neurons from the globus pallidus of parkinsonian primates during passive limb movements and to determine the extent to which they interact and synchronize. The vast majority (80/103) of the recorded pallidal neurons responded to periodic flexion-extension movements of the elbow. The response pattern was sinusoidal-like and the timing of the peak response of the neurons was uniformly distributed around the movement cycle. The interaction between the neuronal activities was analyzed for 123 simultaneously recorded pairs of neurons. Movement-based signal correlation values were diverse and their mean was not significantly different from zero, demonstrating that the neurons were not activated synchronously in response to movement. Additionally, the difference in the peak responses phase of pairs of neurons was uniformly distributed, showing their independent firing relative to the movement cycle. Our results indicate that despite the widely distributed activity in the globus pallidus of the parkinsonian primate, movement encoding is dispersed and independent rather than correlated and synchronized, thus contradicting current views that posit synchronous activation during Parkinson's disease.     

Neuronal activity of the external oculomotor nucleus during saccadic eye movement in the waking cat

The activity of antidromically identified abducens nucleus motoneurons and inter-nuclear neurons has been recorded during saccadic eye movements in the alert cat. The activity of these neurons has been demonstrated to be the sum of a velocity component proportional to eye velocity plus a position component proportional to instantaneous eye position during the movement. Results are discussed in relation to proposed models about the generation of saccadic eye movements.     

Do bimanual motor actions involve the dorsal premotor (PMd), cingulate (CMA) and posterior parietal (PPC) cortices? Comparison with primary and supplementary motor cortical areas

Single neuronal activity was recorded from the dorsal premotor cortex (PMd), the cingulate motor area (CMA) and the posterior parietal cortex (PPC) in two Macaca fascicularis trained to perform a delayed conditional sequence of coordinated pull and grasp movements. The monkey had to perform three types of trials instructed in a random manner: (i) bimanually, using the two hands in a coordinated sequence of movements; (ii) unimanually, using the left hand only; (iii) unimanually, using the right hand only. The aim of this study was first to assess the bilateral relationships of the three cortical areas for unimanual motor control. Second, to establish whether the three cortical areas contain units reflecting bimanual synergy. A total of 255 task-related neurons were recorded from the PMd, CMA and PPC, where most neurons exhibited a significant modulation of activity in both contralateral and ipsilateral unimanual trials (bilateral neurons: 85, 77 and 61%, respectively). Lower proportions of neurons in PMd (7%), CMA (16%) and PPC (6%) were active in unimanual contralateral trials, but not in unimanual ipsilateral trials. The reverse (modulation of activity in ipsilateral but not contralateral unimanual trials) represented 5% of neurons in PMd, 7% in CMA and 3% in PPC. When comparing unimanual and bimanual trials to search evidence for bimanual coordination, 57% of PMd task-related neurons were classified as bimanual, defined as units in which the activity observed in bimanual trials could not be predicted from that associated with unimanual trials when comparing the same events related to the same arm. The proportion of bimanual neurons in CMA (56%) was comparable to that found in PMd (55%), whereas PPC exhibited a higher proportion of bimanual neurons (74%). Furthermore, comparison of the present data with our previous results regarding the supplementary (SMA) and primary (M1) motor cortical areas shows that there is no statistically significant difference between PMd, CMA, SMA and M1 with respect to the proportions of bimanual neurons. Altogether, these results suggest that the five cortical areas PMd, CMA, PPC, SMA and M1 are participating to the control of sequential bimanually coordinated movements. Inter-limb coordination may thus be controlled by a widely distributed network including several cortical and sub-cortical areas.     

Do bimanual motor actions involve the dorsal premotor (PMd), cingulate (CMA) and posterior parietal (PPC) cortices? Comparison with primary and supplementary motor cortical areas

Single neuronal activity was recorded from the dorsal premotor cortex (PMd), the cingulate motor area (CMA) and the posterior parietal cortex (PPC) in two Macaca fascicularis trained to perform a delayed conditional sequence of coordinated pull and grasp movements. The monkey had to perform three types of trials instructed in a random manner: (i) bimanually, using the two hands in a coordinated sequence of movements; (ii) unimanually, using the left hand only; (iii) unimanually, using the right hand only. The aim of this study was first to assess the bilateral relationships of the three cortical areas for unimanual motor control. Second, to establish whether the three cortical areas contain units reflecting bimanual synergy. A total of 255 task-related neurons were recorded from the PMd, CMA and PPC, where most neurons exhibited a significant modulation of activity in both contralateral and ipsilateral unimanual trials (bilateral neurons: 85, 77 and 61%, respectively). Lower proportions of neurons in PMd (7%), CMA (16%) and PPC (6%) were active in unimanual contralateral trials, but not in unimanual ipsilateral trials. The reverse (modulation of activity in ipsilateral but not contralateral unimanual trials) represented 5% of neurons in PMd, 7% in CMA and 3% in PPC. When comparing unimanual and bimanual trials to search evidence for bimanual coordination, 57% of PMd task-related neurons were classified as bimanual, defined as units in which the activity observed in bimanual trials could not be predicted from that associated with unimanual trials when comparing the same events related to the same arm. The proportion of bimanual neurons in CMA (56%) was comparable to that found in PMd (55%), whereas PPC exhibited a higher proportion of bimanual neurons (74%). Furthermore, comparison of the present data with our previous results regarding the supplementary (SMA) and primary (M1) motor cortical areas shows that there is no statistically significant difference between PMd, CMA, SMA and M1 with respect to the proportions of bimanual neurons. Altogether, these results suggest that the five cortical areas PMd, CMA, PPC, SMA and M1 are participating to the control of sequential bimanually coordinated movements. Inter-limb coordination may thus be controlled by a widely distributed network including several cortical and sub-cortical areas.     

Neuronal responses in area 7 of the cat cortex during defensive conditioning

Responses of neurons in area 7 of the parietal association cortex during and after formation of a defensive conditioned reflex to sound were recorded in waking cats. Changes in spike responses of the neurons as a result of the onset of conditioned reflex limb movements were observed in 68% of neurons. Spike responses of neurons formed as a result of learning appeared only if conditioned-reflex limb movements appeared, and they were not observed if, for some reason or other, movements were absent after presentation of the positive conditioned stimulus or on extinction of the reflex. Responses of 46% neurons to conditioned stimulation preceded the conditioned-reflex motor responses by 50–450 msec. The remaining responding neurons were recruited into the response after the beginning of movement. Characteristic spike responses of neurons to the conditioned stimulus appeared 500–900 msec before the beginning of movement and, in the case of appearance of special, "prolonged" motor responses of limb withdrawal, evoked by subsequent reinforcing stimulation.     

Neuronal activity in the primate supplementary motor area and the primary motor cortex in relation to spatio-temporal bimanual coordination

Single neuronal activity was recorded from the supplementary motor area (SMA-proper and pre-SMA) and primary motor cortex (M1) in two Macaca fascicularis trained to perform a delayed conditional sequence of coordinated bimanual pull and grasp movements. The behavioural paradigm was designed to distinguish neuronal activity associated with bimanual coordination from that related to a comparable motor sequence but executed unimanually (left or right arm only). The bimanual and unimanual trials were instructed in a random order by a visual cue. Following the cue, there was a waiting period until presentation of a "go-signal", signalling the monkey to perform the instructed movement. A total of 143 task-related neurons were recorded from the SMA (SMA-proper, 62; pre-SMA, 81). Most SMA units (87%) were active in both unimanual contralateral and unimanual ipsilateral trials (bilateral neurons), whereas 9% of units were active only in unimanual contralateral trials and 3% were active only in unimanual ipsilateral trials. Forty-eight per cent of SMA task-related units were classified as bimanual, defined as neurons in which the activity observed in bimanual trials could not be predicted from that associated with unimanual trials when comparing the same events related to the same arm. For direct comparison, 527 neurons were recorded from M1 in the same monkeys performing the same tasks. The comparison showed that M1 contains significantly less bilateral neurons (75%) than the SMA, whereas the reverse was observed for contralateral neurons (22% in M1). The proportion of M1 bimanual cells (53%) was not statistically different from that observed in the SMA. The results suggest that both the SMA and M1 may contribute to the control of sequential bimanual coordinated movements. Interlimb coordination may then take place in a distributed network including at least the SMA and M1, but the contribution of other cortical and subcortical areas such as cingulate motor cortex and basal ganglia remains to be investigated.     

Correlation of unit activity of the right and left amygdala in food motivation and emotional stress

Correlation between activities of neurons in the right and left central nuclei of amygdala of rabbits recorded during quiet wakefulness, after 24-h food deprivation, after satiation and during emotional stress (demonstration of a dog) was studied by plotting crosscorrelation histograms. The histogram peaks shifted from zero were observed in 50-67% cases. In hungry animals, in a greater number of cases (52%), the discharge of a neuron in the left amygdala was the first in a pair, and the discharge of the right neuron was delayed (peaks from 10 to 50 and from 130 to 150 ms). The opposite order of discharges was less frequent (36%). When a rabbit saw a dog, the number of common inputs to neurons increased and the leading role of the right amygdalar neurons grew (57%) due to an increase in inhibitory influences from the right to the left amygdala. In most cases, the interaction between amygdalar neurons occurred at the frequencies of the delta range, predominantly, from 2 to 4 Hz.     

Seasonal occurrence and local movements of the grey-headed (brown-necked) parrot Poicephalus fuscicollis suahelicus in southern Africa

Seasonal movements of grey‐headed (brown‐necked) parrots were recorded in parts of its range and are likely a response to breeding and availability of specific food sources. Breeding occurred in the northern Kruger National Park and lowveld near the Mutale–Luvhuvhu river confluence from April to August. Aggregations and movements of birds occurred during the post‐breeding season (August–December) in response to seasonally abundant food sources. In north‐eastern South Africa, grey‐headed parrots occurred at Levubu, following the breeding season and their arrival in the area was correlated with the availability of unripe Mabola Plum, Parinari curatellifolia fruit. Similar regional movements occurred in Zimbabwe, the Caprivi of northern Namibia and Zambia. During these movements, flocks of up to 50 individuals were observed, whilst during breeding months singletons and pairs were more frequently seen. This increased abundance in time and space suggests that seasonal migratory movements occur.     

Control of jaw movements in two species of macropodines (Macropus eugenii and Macropus rufus)

The masticatory motor patterns of three tammar wallabies and two red kangaroos were determined by analyzing the pattern of electromyographic (EMG) activity of the jaw adductors and correlating it with lower jaw movements, as recorded by digital video and videoradiography. Transverse jaw movements were limited by the width of the upper incisal arcade. Molars engaged in food breakdown during two distinct occlusal phases characterized by abrupt changes in the direction of working-side hemimandible movement. Separate orthal (Phase I) and transverse (Phase II) trajectories were observed. The working-side lower jaw initially was drawn laterally by the balancing-side medial pterygoid and then orthally by overlapping activity in the balancing- and working-side temporalis and the balancing-side superficial masseter and medial pterygoid. Transverse movement occurred principally via the working-side medial pterygoid and superficial masseter. This pattern contrasted to that of placental herbivores, which are known to break down food when they move the working-side lower jaw transversely along a relatively longer linear path without changing direction during the power stroke. The placental trajectory results from overlapping activity in the working- and balancing-side adductor muscles, suggesting that macropods and placental herbivores have modified the primitive masticatory motor pattern in different ways.