Functional mapping of the circuits involved in the expression of contextual fear responses in socially defeated animals.

In this study, we have aimed at outlining the neural systems underlying the expression of contextual fear to social defeat. First, we have developed an experimental procedure, where defeated animals could express, without the presence of a dominant aggressive male, robust and reliable conditioned fear responses to the context associated with social defeat. Next, by examining the pattern of Fos expression, we have been able to outline a brain circuit comprising septal and amygdalar sites, as well as downstream hypothalamic paths, putatively involved in the expression of contextual fear to social threat. Of particular relevance, we have found that exposure to a defeat-associated context results in a striking Fos up-regulation in the dorsomedial part of the dorsal premammillary nucleus (PMDdm). To further understand the role of the PMDdm in the circuit organizing conditioned fear to social threats, we have been able to observe that pharmacological blockade of the PMDdm reduced fear responses to a social defeat-associated context. Next, we observed that pharmacological blockade of the dorsomedial part of the periaqueductal gray, one of the main targets of the PMDdm, produced an even higher reduction of conditioned fear in defeated intruders, and appears as an important node for the expression of contextual defensive responses to social threats. The present results help to elucidate the basic organization of the neural circuits underlying contextual conditioned responses to social defeat, and reveal that they share at least part of the same circuit involved in innate responses to social defeat to an aggressive conspecific.

Experts in offside decision making learn to compensate for their illusory perceptions.

In association football, the flash-lag effect appears to be a viable explanation for erroneous offside decision making. Due to this spatiotemporal illusion, assistant referees (ARs) perceive the player who receives the ball ahead of his real position. In this experiment, a laboratory decision-making task was used to demonstrate that international top-class ARs, compared with amateur soccer players, do not have superior perceptual sensitivity. They clearly modify their decision criterion according to the contextual needs and, therefore, show a higher response bias toward not responding to the stimulus, in particular in the most difficult situations. Thus, international ARs show evidence for response-level compensation, resulting in a specific cost (i.e., more misses), which clearly reflects the use of particular (cognitive) strategies. In summary, it appears that experts in offside decision making can be distinguished from novices more on the cognitive or decision-making level than on the perceptual level.

Amygdalar roles during exposure to a live predator and to a predator-associated context.

The amygdala plays a critical role in determining the emotional significance of sensory stimuli and the production of fear-related responses. Large amygdalar lesions have been shown to practically abolish innate defensiveness to a predator; however, it is not clear how the different amygdalar systems participate in the defensive response to a live predator. Our first aim was to provide a comprehensive analysis of the amygdalar activation pattern during exposure to a live cat and to a predator-associated context. Accordingly, exposure to a live predator up-regulated Fos expression in the medial amygdalar nucleus (MEA) and in the lateral and posterior basomedial nuclei, the former responding to predator-related pheromonal information and the latter two nuclei likely to integrate a wider array of predatory sensory information, ranging from olfactory to non-olfactory ones, such as visual and auditory sensory inputs. Next, we tested how the amygdalar nuclei most responsive to predator exposure (i.e. the medial, posterior basomedial and lateral amygdalar nuclei) and the central amygdalar nucleus (CEA) influence both unconditioned and contextual conditioned anti-predatory defensive behavior. Medial amygdalar nucleus lesions practically abolished defensive responses during cat exposure, whereas lesions of the posterior basomedial or lateral amygdalar nuclei reduced freezing and increased risk assessment displays (i.e. crouch sniff and stretch postures), a pattern of responses compatible with decreased defensiveness to predator stimuli. Moreover, the present findings suggest a role for the posterior basomedial and lateral amygdalar nuclei in the conditioning responses to a predator-related context. We have further shown that the CEA does not seem to be involved in either unconditioned or contextual conditioned anti-predatory responses. Overall, the present results help to clarify the amygdalar systems involved in processing predator-related sensory stimuli and how they influence the expression of unconditioned and contextual conditioned anti-predatory responses.

The role of the superior colliculus in predatory hunting.

Combining the results of behavioral, neuronal immediate early gene activation, lesion and neuroanatomical experiments, we have presently investigated the role of the superior colliculus (SC) in predatory hunting. First, we have shown that insect hunting is associated with a characteristic large increase in Fos expression in the lateral part of the intermediate gray layer of the SC (SCig). Next, we have shown that animals with bilateral NMDA lesions of the lateral parts of the SC presented a significant delay in starting to chase the prey and longer periods engaged in other activities than predatory hunting. They also showed a clear deficit to orient themselves toward the moving prey and lost the stereotyped sequence of actions seen for capturing, holding and killing the prey. Our Phaseolus vulgaris-leucoagglutinin analysis revealed that the lateral SCig, besides providing the well-documented descending crossed pathway to premotor sites in brainstem and spinal cord, projects to a number of midbrain and diencephalic sites likely to influence key functions in the context of the predatory behavior, such as general levels of arousal, motivational level to hunt or forage, behavioral planning, appropriate selection of the basal ganglia motor plan to hunt, and motor output of the primary motor cortex. In contrast to the lateral SC lesions, medial SC lesions produced a small deficit in predatory hunting, and compared to what we have seen for the lateral SCig, the medial SCig has a very limited set of projections to thalamic sites related to the control of motor planning or motor output, and provides conspicuous inputs to brainstem sites involved in organizing a wide range of anti-predatory defensive responses. Overall, the present results served to clarify how the different functional domains in the SC may mediate the decision to pursue and hunt a prey or escape from a predator.

Hypothalamic sites responding to predator threats--the role of the dorsal premammillary nucleus in unconditioned and conditioned antipredatory defensive behavior.

In this study we provide a comprehensive analysis of the hypothalamic activation pattern during exposure to a live predator or an environment previously associated with a predator. Our results support the view that hypothalamic processing of the actual and the contextual predatory threats share the same circuit, in which the dorsal premammillary nucleus (PMd) plays a pivotal role in amplifying this processing. To further understand the role of the PMd in the circuit organizing antipredatory defensive behaviors, we studied rats with cytotoxic PMd lesions during cat exposure and examined the pattern of behavioral responses as well as how PMd lesions affect the neuronal activation of the systems engaged in predator detection, in contextual memory formation and in defensive behavioral responses. Next, we investigated how pharmacological blockade of the PMd interferes with the conditioned behavioral responses to a context previously associated with a predator, and how this blockade affects the activation pattern of periaqueductal gray (PAG) sites likely to organize the conditioned behavioral responses to the predatory context. Behavioral observations indicate that the PMd interferes with both unconditioned and conditioned antipredatory defensive behavior. Moreover, we have shown that the PMd influences the activation of its major projecting targets, i.e. the ventral part of the anteromedial thalamic nucleus which is likely to influence mnemonic processing, and PAG sites involved in the expression of antipredatory unconditioned and conditioned behavioral responses. Of particular relevance, this work provides evidence to elucidate the basic organization of the neural circuits integrating unconditioned and contextual conditioned responses to predatory threats.

A role for the periaqueductal gray in switching adaptive behavioral responses.

Previous studies suggested a role for the rostral lateral periaqueductal gray (PAG) in the inhibition of maternal behavior induced by low doses of morphine in dams with previous morphine experience. In the present study, we first showed that unilateral NMDA lesions placed in this particular PAG region prevented the morphine-induced inhibition of maternal behavior in previously morphine-sensitized dams. As suggested by previous Fos data on the PAG, predatory hunting appears as a likely candidate to replace maternal behavior in the morphine-treated dams. By testing saline- and morphine-treated dams with live cockroaches only, we have presently shown that morphine challenge increased insect hunting. Moreover, morphine- and saline-treated dams were also observed in an environment containing pups and roaches. Although most of the saline-treated animals displayed active nursing and only occasionally presented insect hunting, all of the morphine-treated animals ignored the pups and avidly pursued and caught the roaches. We next questioned whether the rostral lateral PAG would be involved in this behavioral switch. Our results showed that unilateral lesions of the rostral lateral PAG, but not other parts of the PAG, partially impaired predatory hunting and restored part of the maternal response. Moreover, bilateral lesions of the rostral lateral PAG produced even more dramatic effects in inhibiting insect hunting and restoring maternal behavior. The present findings indisputably show that the rostral lateral PAG influences switching from maternal to hunting behavior in morphine-treated dams, thus supporting a previously unsuspected role for the PAG in selecting adaptive behavioral responses.

The decreased oxygen uptake during progressive exercise in ischemia-induced heart failure is due to reduced cardiac output rate.

We tested the hypothesis that the inability to increase cardiac output during exercise would explain the decreased rate of oxygen uptake (VO2) in recent onset, ischemia-induced heart failure rats. Nine normal control rats and 6 rats with ischemic heart failure were studied. Myocardial infarction was induced by coronary ligation. VO2 was measured during a ramp protocol test on a treadmill using a metabolic mask. Cardiac output was measured with a flow probe placed around the ascending aorta. Left ventricular end-diastolic pressure was higher in ischemic heart failure rats compared with normal control rats (17 +/- 0.4 vs 8 +/- 0.8 mmHg, P = 0.0001). Resting cardiac index (CI) tended to be lower in ischemic heart failure rats (P = 0.07). Resting heart rate (HR) and stroke volume index (SVI) did not differ significantly between ischemic heart failure rats and normal control rats. Peak VO2 was lower in ischemic heart failure rats (73.72 +/- 7.37 vs 109.02 +/- 27.87 mL min(-1) kg(-1), P = 0.005). The VO2 and CI responses during exercise were significantly lower in ischemic heart failure rats than in normal control rats. The temporal response of SVI, but not of HR, was significantly lower in ischemic heart failure rats than in normal control rats. Peak CI, HR, and SVI were lower in ischemic heart failure rats. The reduction in VO2 response during incremental exercise in an ischemic model of heart failure is due to the decreased cardiac output response, largely caused by depressed stroke volume kinetics.

The decreased oxygen uptake during progressive exercise in ischemia-induced heart failure is due to reduced cardiac output rate

We tested the hypothesis that the inability to increase cardiac output during exercise would explain the decreased rate of oxygen uptake (VO2) in recent onset, ischemia-induced heart failure rats. Nine normal control rats and 6 rats with ischemic heart failure were studied. Myocardial infarction was induced by coronary ligation. VO2 was measured during a ramp protocol test on a treadmill using a metabolic mask. Cardiac output was measured with a flow probe placed around the ascending aorta. Left ventricular end-diastolic pressure was higher in ischemic heart failure rats compared with normal control rats (17 ± 0.4 vs 8 ± 0.8 mmHg, P = 0.0001). Resting cardiac index (CI) tended to be lower in ischemic heart failure rats (P = 0.07). Resting heart rate (HR) and stroke volume index (SVI) did not differ significantly between ischemic heart failure rats and normal control rats. Peak VO2 was lower in ischemic heart failure rats (73.72 ± 7.37 vs 109.02 ± 27.87 mL min-1 kg-1, P = 0.005). The VO2 and CI responses during exercise were significantly lower in ischemic heart failure rats than in normal control rats. The temporal response of SVI, but not of HR, was significantly lower in ischemic heart failure rats than in normal control rats. Peak CI, HR, and SVI were lower in ischemic heart failure rats. The reduction in VO2 response during incremental exercise in an ischemic model of heart failure is due to the decreased cardiac output response, largely caused by depressed stroke volume kinetics.

The modulation of simple reaction time by the spatial probability of a visual stimulus.

Simple reaction time (SRT) in response to visual stimuli can be influenced by many stimulus features. The speed and accuracy with which observers respond to a visual stimulus may be improved by prior knowledge about the stimulus location, which can be obtained by manipulating the spatial probability of the stimulus. However, when higher spatial probability is achieved by holding constant the stimulus location throughout successive trials, the resulting improvement in performance can also be due to local sensory facilitation caused by the recurrent spatial location of a visual target (position priming). The main objective of the present investigation was to quantitatively evaluate the modulation of SRT by the spatial probability structure of a visual stimulus. In two experiments the volunteers had to respond as quickly as possible to the visual target presented on a computer screen by pressing an optic key with the index finger of the dominant hand. Experiment 1 (N = 14) investigated how SRT changed as a function of both the different levels of spatial probability and the subject's explicit knowledge about the precise probability structure of visual stimulation. We found a gradual decrease in SRT with increasing spatial probability of a visual target regardless of the observer's previous knowledge concerning the spatial probability of the stimulus. Error rates, below 2%, were independent of the spatial probability structure of the visual stimulus, suggesting the absence of a speed-accuracy trade-off. Experiment 2 (N = 12) examined whether changes in SRT in response to a spatially recurrent visual target might be accounted for simply by sensory and temporally local facilitation. The findings indicated that the decrease in SRT brought about by a spatially recurrent target was associated with its spatial predictability, and could not be accounted for solely in terms of sensory priming.

The modulation of simple reaction time by the spatial probability of a visual stimulus

Simple reaction time (SRT) in response to visual stimuli can be influenced by many stimulus features. The speed and accuracy with which observers respond to a visual stimulus may be improved by prior knowledge about the stimulus location, which can be obtained by manipulating the spatial probability of the stimulus. However, when higher spatial probability is achieved by holding constant the stimulus location throughout successive trials, the resulting improvement in performance can also be due to local sensory facilitation caused by the recurrent spatial location of a visual target (position priming). The main objective of the present investigation was to quantitatively evaluate the modulation of SRT by the spatial probability structure of a visual stimulus. In two experiments the volunteers had to respond as quickly as possible to the visual target presented on a computer screen by pressing an optic key with the index finger of the dominant hand. Experiment 1 (N = 14) investigated how SRT changed as a function of both the different levels of spatial probability and the subject's explicit knowledge about the precise probability structure of visual stimulation. We found a gradual decrease in SRT with increasing spatial probability of a visual target regardless of the observer's previous knowledge concerning the spatial probability of the stimulus. Error rates, below 2 percent, were independent of the spatial probability structure of the visual stimulus, suggesting the absence of a speed-accuracy trade-off. Experiment 2 (N = 12) examined whether changes in SRT in response to a spatially recurrent visual target might be accounted for simply by sensory and temporally local facilitation. The findings indicated that the decrease in SRT brought about by a spatially recurrent target was associated with its spatial predictability, and could not be accounted for solely in terms of sensory priming

The attentional modulation of the flash-lag effect.

If a dot is flashed in perfect alignment with a pair of dots rotating around the visual fixation point, most observers perceive the rotating dots as being ahead of the flashing dot (flash-lag effect). This perceptual effect has been interpreted to result from the perceptual extrapolation of the moving dots, the differential visual latencies between flashing and moving stimuli, as well as the modulation of attentional mechanisms. Here we attempted to uncouple the attentional effects brought about by the spatial predictability of the flashing dot from the sensory effects dependent on its visual eccentricity. The stimulus was a pair of dots rotating clockwise around the fixation point. Another dot was flashed at either the upper right or the lower left of the visual field according to three separate blocked situations: fixed, alternate and random positions. Twenty-four participants had to judge, in all three situations, the location of the rotating dots in relation to the imaginary line connecting the flashing dot and the fixation point at the moment the dot was flashed. The flash-lag effect was observed in all three situations, and a clear influence of the spatial predictability of the flashing dot on the magnitude of the perceptual phenomenon was revealed, independently of sensory effects related to the eccentricity of the stimulus in the visual field. These findings are consistent with our proposal that, in addition to sensory factors, the attentional set modulates the magnitude of the differential latencies that give rise to the flash-lag phenomenon.

Modulation of the perception of temporal order by attentional and pre-attentional factors.

When two stimuli are presented simultaneously to an observer, the perceived temporal order does not always correspond to the actual one. In three experiments we examined how the location and spatial predictability of visual stimuli modulate the perception of temporal order. Thirty-two participants had to report the temporal order of appearance of two visual stimuli. In Experiment 1, both stimuli were presented at the same eccentricity and no perceptual asynchrony between them was found. In Experiment 2, one stimulus was presented close to the fixation point and the other, peripheral, stimulus was presented in separate blocks in two eccentricities (4.8 degrees and 9.6 degrees ). We found that the peripheral stimulus was perceived to be delayed in relation to the central one, with no significant difference between the delays obtained in the two eccentricities. In Experiment 3, using three eccentricities (2.5 degrees, 7.3 degrees and 12.1 degrees ) for the presentation of the peripheral stimulus, we compared a condition in which its location was highly predictable with two other conditions in which its location was progressively less predictable. Here, the perception of the peripheral stimulus was also delayed in relation to the central one, with this delay depending on both the eccentricity and predictability of the stimulus. We argue that attentional deployment, manipulated by the spatial predictability of the stimulus, seems to play an important role in the temporal order perception of visual stimuli. Yet, under whichever condition of spatial predictability, basic sensory and attentional processes are unavoidably entangled and both factors must concur to the perception of temporal order.

Modulation of the perception of temporal order by attentional and pre-attentional factors

When two stimuli are presented simultaneously to an observer, the perceived temporal order does not always correspond to the actual one. In three experiments we examined how the location and spatial predictability of visual stimuli modulate the perception of temporal order. Thirty-two participants had to report the temporal order of appearance of two visual stimuli. In Experiment 1, both stimuli were presented at the same eccentricity and no perceptual asynchrony between them was found. In Experiment 2, one stimulus was presented close to the fixation point and the other, peripheral, stimulus was presented in separate blocks in two eccentricities (4.8º and 9.6º). We found that the peripheral stimulus was perceived to be delayed in relation to the central one, with no significant difference between the delays obtained in the two eccentricities. In Experiment 3, using three eccentricities (2.5º, 7.3º and 12.1º) for the presentation of the peripheral stimulus, we compared a condition in which its location was highly predictable with two other conditions in which its location was progressively less predictable. Here, the perception of the peripheral stimulus was also delayed in relation to the central one, with this delay depending on both the eccentricity and predictability of the stimulus. We argue that attentional deployment, manipulated by the spatial predictability of the stimulus, seems to play an important role in the temporal order perception of visual stimuli. Yet, under whichever condition of spatial predictability, basic sensory and attentional processes are unavoidably entangled and both factors must concur to the perception of temporal order

The attentional modulation of the flash-lag effect

If a dot is flashed in perfect alignment with a pair of dots rotating around the visual fixation point, most observers perceive the rotating dots as being ahead of the flashing dot (flash-lag effect). This perceptual effect has been interpreted to result from the perceptual extrapolation of the moving dots, the differential visual latencies between flashing and moving stimuli, as well as the modulation of attentional mechanisms. Here we attempted to uncouple the attentional effects brought about by the spatial predictability of the flashing dot from the sensory effects dependent on its visual eccentricity. The stimulus was a pair of dots rotating clockwise around the fixation point. Another dot was flashed at either the upper right or the lower left of the visual field according to three separate blocked situations: fixed, alternate and random positions. Twenty-four participants had to judge, in all three situations, the location of the rotating dots in relation to the imaginary line connecting the flashing dot and the fixation point at the moment the dot was flashed. The flash-lag effect was observed in all three situations, and a clear influence of the spatial predictability of the flashing dot on the magnitude of the perceptual phenomenon was revealed, independently of sensory effects related to the eccentricity of the stimulus in the visual field. These findings are consistent with our proposal that, in addition to sensory factors, the attentional set modulates the magnitude of the differential latencies that give rise to the flash-lag phenomenon

Accessory optic-pretectal interactions in the pigeon.

This electrophysiological study analyzes the influence of the nucleus of the basal optic root (nBOR) of the avian accessory optic system on units within the lentiform nucleus (LM), which is the avian equivalent of the pretectal nucleus of the optic tract. A prominent depression of the spontaneous firing rate of neurons within the LM occurred following electrical stimulation of the ipsilateral nBOR. A close correlation was also found between the directional selectivity of LM units and the apparent displacements generated by rotations of the head around the horizontal semicircular canal axis. This is consistent with a possible role of the LM in the coordinate transformation from visual inputs to a vestibular reference system.

The spatial arrangement of the semicircular canals of the pigeon.

The spatial orientation of the semicircular canals (SCC) was measured in the pigeon by fitting the best plane over 10 to 15 stereotaxically determined points to each canal. The vectors defining each canal plane are presented in the standard stereotaxic position and for free moving behavior, and the coplanarity or orthogonality of pairs of semicircular canals are discussed.

Accessory optic-pretectal interactions in the pigeon

This electrophysiological study analyzes the influence of the nucleus of the basal optic root (nBOR) of the avian accessory optic sustem on units within the lentiform nucleus (LM), which is the avian equivalent of the pretectal nucleus of the optic trat. A prominent depression of the spontaneous firing rat of neurons within the LM occurred following electrical stimulation of the ipsilateral nBOR. A close correlation was also found between the directional selectivity of LM units and the apparent displacements generated by rotations of the head around the horizontal semicircular canal axis. This is consistent with a possible rolo of the LM in the coordinate transformation from visual imputs to a vestibular referense system

The spatial arrangement of the semicircular canals of the pigeon

The spatial orientation of the semicircular canals (SCC) was measured in the pigeon by fitting the best plane over 10 to 15 stereotaxically determined points to each canal. The vectores definign eah canal plane are presented in the standard stereotaxical position and for free moving behavior, and the coplanarity or orthogonality of pairs of semicircular canals are discussed

The relation between head and apparent visual field motion in the pigeon(Columba livia)

In order to study the relationship the apparent motion of the visual field as a function of head motion, we have developed an equation that describes the apparent displacement of objects in the visual field produced by a general movement of the head consisting of a rotation plus a translation. We then used this equation to simulate the apparent motion of the visual field of a pigeon produced by rotation of the head about axes parallel to those of its anterior semicircular canals