T-pattern analysis for the study of temporal structure of animal and human behavior: a comprehensive review.

A basic tenet in the realm of modern behavioral sciences is that behavior consists of patterns in time. For this reason, investigations of behavior deal with sequences that are not easily perceivable by the unaided observer. This problem calls for improved means of detection, data handling and analysis. This review focuses on the analysis of the temporal structure of behavior carried out by means of a multivariate approach known as T-pattern analysis. Using this technique, recurring sequences of behavioral events, usually hard to detect, can be unveiled and carefully described. T-pattern analysis has been successfully applied in the study of various aspects of human or animal behavior such as behavioral modifications in neuro-psychiatric diseases, route-tracing stereotypy in mice, interaction between human subjects and animal or artificial agents, hormonal-behavioral interactions, patterns of behavior associated with emesis and, in our laboratories, exploration and anxiety-related behaviors in rodents. After describing the theory and concepts of T-pattern analysis, this review will focus on the application of the analysis to the study of the temporal characteristics of behavior in different species from rodents to human beings. This work could represent a useful background for researchers who intend to employ such a refined multivariate approach to the study of behavior.

Multivariate temporal pattern analysis applied to the study of rat behavior in the elevated plus maze: methodological and conceptual highlights.

Aim of this article is to illustrate the application of a multivariate approach known as t-pattern analysis in the study of rat behavior in elevated plus maze. By means of this multivariate approach, significant relationships among behavioral events in the course of time can be described. Both quantitative and t-pattern analyses were utilized to analyze data obtained from fifteen male Wistar rats following a trial 1-trial 2 protocol. In trial 2, in comparison with the initial exposure, mean occurrences of behavioral elements performed in protected zones of the maze showed a significant increase counterbalanced by a significant decrease of mean occurrences of behavioral elements in unprotected zones. Multivariate t-pattern analysis, in trial 1, revealed the presence of 134 t-patterns of different composition. In trial 2, the temporal structure of behavior become more simple, being present only 32 different t-patterns. Behavioral strings and stripes (i.e. graphical representation of each t-pattern onset) of all t-patterns were presented both for trial 1 and trial 2 as well. Finally, percent distributions in the three zones of the maze show a clear-cut increase of t-patterns in closed arm and a significant reduction in the remaining zones. Results show that previous experience deeply modifies the temporal structure of rat behavior in the elevated plus maze. In addition, this article, by highlighting several conceptual, methodological and illustrative aspects on the utilization of t-pattern analysis, could represent a useful background to employ such a refined approach in the study of rat behavior in elevated plus maze.

Significant divergences between the temporal structure of the behavior in Wistar and in the spontaneously more anxious DA/Han strain of rats tested in elevated plus maze.

The aim of present research is to study the temporal structure of the behavior in two strains of rats with different basal level of emotionality. To this purpose, the temporal profile of the behavior in Wistar rat and in the spontaneously more anxious DA/Han strain was analyzed in the Elevated Plus Maze. Both quantitative and multivariate t-pattern analyses were carried out. In comparison with Wistar, DA/Han subjects showed a significant reduction of the permanence in open arm and a significant increase of the time spent in the central platform of the maze. Mean frequencies of each behavioral element showed significant modifications both in open and in closed arm. Multivariate t-pattern analyses demonstrated a very different temporal profile of behavior in the two strains: Wistar rats presented 197 t-patterns of different composition, whereas DA/Han rats only 26; as to the mean number of t-patterns, Wistar presented a value of 698.90, whereas DA/Han only 92.80. Similar clear-cut differences were detected for the mean number of t-patterns in the two arms and for the time course of such t-patterns. Present study has evidenced that the temporal organization of the bahavior in Elevated Plus Maze is differently structured in two strains of rats with different basal level of emotionality.

Temporal structure of the rat's behavior in elevated plus maze test.

Aim of the research was to evaluate, by means of quantitative and multivariate temporal pattern analyses, the behavior of Wistar rat in elevated plus maze (EPM) test. On the basis of an ethogram encompassing 24 behavioral elements, quantitative results showed that 130.14 ± 8.01 behavioral elements occurred in central platform and in closed arms (protected zones), whereas 88.62 ± 6.04 occurred in open arms (unprotected zones). Percent distribution was characterized by a prevalence of sniffing, walking and vertical exploration. Analysis of minute-by-minute duration evidenced a decrease for time spent in open arms and central platform and an increase for time spent in closed arms. As to multivariate t-pattern analysis, 126 different temporal patterns were detected. Behavioral stripes, summarizing distribution of such t-patterns along time, showed that several t-patterns were not homogeneously distributed along the test observational period: t-patterns encompassing behavioral events occurring prevalently in central platform-open arms were observed during the first minutes, whereas t-patterns structured on the basis of events occurring mainly in central platform-closed arms were detected during the last minutes. Therefore, during the observation in elevated plus maze, rat's behavior undergoes significant rearrangements of its temporal features. Present research demonstrates, for the first time, the existence of complex and significantly timed behavioral sequences in the activity of Wistar rats tested in elevated plus maze. Application of t-pattern analysis can provide useful tools to characterize the behavioral dynamics of anxiety-related rodent behavior and differentiate the effect of various anxioselective substances.

Inhibitory effect of A10 dopaminergic neurons of the ventral tegmental area on the orienting response evoked by acoustic stimulation in the cat.

The effect of bilateral electric stimulation of A10 dopaminergic neurons of the ventral tegmental area (80-300 microA, 20-50 Hz, 0.1-0.5 ms, 2 s duration) on latency and duration of the orienting response, evoked by acoustic stimuli (4500-8000 Hz, 2 s), was studied in the cat. A10 neuron stimulation, simultaneous with the acoustic one, was performed with threshold parameters inducing minimal behavioral signs (head searching movement, sniffing, increase in alertness). By means of a videoanalysis system, a statistically significant increase, both of latency and duration of the response, was observed. The possible role of dopamine was studied administrating sulpiride (20 mg/kg i.p.), a dopaminergic antagonist prevalently acting on the mesolimbic-mesocortical system. In this condition, the disappearance of A10 neuron effect occurred. Sulpiride injection did not affect the parameters of the orienting response to acoustic stimulus alone, suggesting a direct effect on A10 dopaminergic neurons. Moreover, when saline administration was carried out, no significant modification of the effects, obtained following A10 neuron activation, was observed. The data suggest that A10 dopaminergic neurons, origin of the mesolimbic-mesocortical system, may be involved in the control of the response to sensory stimuli, likely by influencing sensorimotor integration processes. An involvement in the inhibitory regulation of the switching of attention is also discussed.

Influence of ventral tegmental area (A10 region) on flight behaviour elicited by hypothalamic stimulation in the cat.

The influence of the ventral tegmental area (VTA) (dopaminergic A10 group neurons) on flight behaviour, induced by hypothalamic stimulation, was studied in the cat. Co-stimulation of hypothalamus and VTA (ipsi- or contralateral) induced an increase of the flight latency. Slow-motion analysis of flight behaviour showed that this increase was due to the augmentation of the fixation latency (FL) whereas the upright latency (UL) was not modified. Sulpiride injection (50 mg/kg i.p.) provoked the disappearance of VTA effect without affecting the basal behavioural sequence. The results show that DA A10 group neurons increase the attentive component of the flight reaction, suggesting a possible influence of the DA A10 system on the mechanisms modulating focused attention in the animal.

Opiate receptor modifications in the rat brain after chronic treatment with haloperidol and suipiride.

Anatomical, electrophysiological and pharmacological data support the existence of a pronounced interaction between dopamine (DA) and opioids. In particular, chronic administration of DA antagonist drugs modifies opiate peptides and opiate receptors. In this paper we focused, by means of quantitative receptor autoradiography, on the modifications induced by chronic neuroleptic treatment, in patches versus diffuse distribution, of opiate receptors in the striatum, and we also studied the different effects of haloperidol and sulpiride on striatal and cortical receptors. We found a significant decrease of the number of (3H)- naloxone binding sites in the striatal patches of treated animals but no effects in the matrix. We also observed, in haloperidol-treated animals, an increase of (3H)-naloxone binding sites in the medial cortex, and in sulpiride-treated animals an increase in the lateral and dorsal cortex. Two main observations arise from our data: (a) a differential effect is produced by neuroleptic treatment on opiate receptors in patches and in matrix; (b) an opposite influence is exerted by sulpiride and haloperidol on opiate receptors in the cortex and in striatum.

Striatal influence on ingestive behaviour in the cat.

In freely moving cats with chronically implanted electrodes an analysis was made of the effects on feeding behaviour of low-frequency long-duration stimulation of the caudate nucleus, the substantia nigra and the globus pallidus. In all 3 structures a significant reduction of food intake was observed and in the pallidus this reached the point of a complete block of feeding. The effects were always limited to the period of stimulation. At the end of stimulation the animals recovered and took in food quantities equal to those of controls. The results are interpreted on the basis of reciprocal connections between the basal ganglia and the hypothalamus; the role of the striatum on the selection of certain movements and its possible involvement in behaviour is also discussed.

Effects of substantia nigra stimulation on electrical and mechanical activities of the duodeno-jejunal loop, in the cat.

The effect of substantia nigra stimulation on duodeno-jejunal intraluminal pressure and electrical activity was analysed in the cat. An inhibitory influence was noted to occur on both the intestinal pressure and the electrical response activity (ERA): the effect seemed to be in correlation with the stimulus parameters employed. After bilateral vagotomy, the inhibition remained substantially unchanged. The possibility of the existence of a sympathetic pathway mediating the effect is emphasized.

[Feeding behavior of the cat in response to stimulation of the caudate nucleus]. / Comportamento alimentare del gatto in seguito a stimolazione del nucleo cau dato.

In chronic free-moving cats unilateral, low frequency, long duration stimulation of the caudate nucleus causes a significant reduction of food intake. The effects are greatest at the beginning of the stimulation. On cessation of stimulation the animals take enough food to compensate for that lost during the time of stimulation. Results are discussed either considering interrelationships between striatum and lateral hypothalamus, either regarding recent viewpoints distinguishing two striatal compartments: dorsal or "non limbic" striatal region and ventral or "limbic" striatal region.

Changes in latency of evoked potentials after caudate nucleus stimulation.

The activation of the head of the caudate nucleus in cats which have undergone transection of the spinal cord at C1 produces inhibition of evoked potentials in the motor area (pre-cruciate region) and also reduces the peak latency of the negative slow wave. The results of decreased latency are discussed with reference to variations in intensity of the stimulation activating the motor cortex. The role played by the caudate nucleus in the control of the motor system is emphasized.

Feeding behaviour following substantia nigra and globus pallidus stimulation in chronic cats.

It is known that the lesion of the nigrostriatal and pallidofugal systems causes an experimental condition comparable to the hypothalamic sindrome, with severe aphagia and adipsia. It was considered of interest to study the effects of the low frequency long-duration stimulation of the substantia nigra and the globus pallidus on feeding behaviour. Experiments were performed on eight chronic cats with electrodes implanted in SN (pars compacta) and in the GP (pars externa). In the 2 hours of observation the food intake and the number of food-taking acts were calculated. The stimulation was induced continuously in the former hour. The modification in food intake, also on the basis of somatic behavioural observations may be ascribed to motor impairment rather than to involvement of a motivational state.

[Different effects of conditioning stimulation of the caudate nucleus and globus pallidus on the paroxysmal activity of the cat amygdala]. / Differenti effectti della stimolazione condizionante del nucleo caudato e del globo pallido sull'attivita parossistica amigdaloidea nel gatto.

The conditioning stimulation of the globus pallidus both in chronic and acute cats determined a more effective inhibition than caudate nucleus activation on the evoked paroxysmal activity in the amygdala. Lesions by means of kainic acid of the entopeduncular nucleus, has resulted in a significant decrease of the caudate inhibitory effect. The hypothesis we assumed is about a role of the globus pallidus in the control of the evoked amygdaloid after-discharge.

[Effects of conditioning stimulation of the substantia nigra on the epileptic activity of the cat amygdala]. / Effetti della stimolazione condizionante della substantia nigra sull'attivita epilettica amigdaloidea nel gatto.

The conditioning stimulation of the substantia nigra both in chronic and acute cats determined a more effective inhibition than caudate nucleus activation on the evoked paroxysmal activity in the amygdala. Lesions of the substantia nigra by means of kainic acid, produced the disappearance of the inhibition induced by caudate nucleus. An examination of the obtained data and of the literature on this topic, allowed us to identify a possible nigro-amygdaloid direct pathway.

The role of the substantia nigra in the control of amygdaloid paroxysmal activity.

Both in acute and chronic cats focal paroxysmal activity evoked in the ventro-basal complex of the amygdala has been inhibited by substantia nigra conditioning stimulation, to a greater extent, than by caudate nucleus activation. Injection of kainic acid into substantia nigra resulted in the disappearance of the caudate inhibition. It is suggested that the final control, exerted by the striatum on the amygdaloid seizures, occurs by means of the substantia nigra.