The trigeminal sensory relay to reticulospinal neurones in lampreys.

This study was carried out to identify lamprey neurones relaying trigeminal sensory inputs to reticulospinal cells. Double labeling with fluorescent tracers was used in vitro. Fluorescein-conjugated dextran amines were applied to the proximal stump of the cut trigeminal nerve on both sides, and Texas Red-conjugated dextran amines were injected unilaterally in the middle (MRRN) or the posterior (PRRN) rhombencephalic reticular nuclei. Texas Red retrogradely labeled cells were found ipsi- and contralateral to each injection. Any of these cells with the soma or at least a major dendrite among the fluorescein-labeled trigeminal afferent axons was considered a candidate relay cell. Of these two possibilities, only cells with their soma among the fluorescein-labeled trigeminal afferents were found. The candidate relay cells projecting to the MRRN were mostly clustered at the caudal vestibular nerve level within the trigeminal descending tract, whereas the majority of those projecting to the PRRN were located more caudally. The diameter of candidate relay cells ranged from 9.2 to 24.6 mum and 9.2 to 46.1 mum, after MRRN and PRRN injections, respectively. A possible relay function for these cells was tested with electrophysiological experiments. The intracellular responses to trigeminal nerve stimulation were recorded in reticulospinal cells under control conditions and after ejections of a combination of glutamate ionotropic receptor antagonists over the candidate relay cells in small areas along the sulcus limitans. The synaptic responses elicited in MRRN reticulospinal cells were maximally depressed when ejections were made at the level of the vestibular nerve, in accord with the anatomical data. The synaptic responses in PRRN reticulospinal cells showed maximal depression when ejections were made slightly more caudally. Altogether, these results suggest that cells located within the trigeminal descending tract and projecting to reticular nuclei are likely to be the sensory trigeminal relays to reticulospinal neurones in lampreys.

Theta-rhythmically firing neurons in the anterior thalamus: implications for mnemonic functions of Papez's circuit.

In 1937 Papez described an anatomical circuit (or loop) beginning and ending in the hippocampal formation that he proposed subserved emotional experience (Papez, 1937). Specifically, the projections of the circuit were as follows: hippocampal formation--> mammillary bodies--> anterior thalamus--> cingulate cortex--> parahippocampal gyrus--> hippocampal formation. Although the circuit has been refined based on subsequent anatomical findings (Amaral and Witter, 1995; Shibata, 1992; Van Groen and Wyss, 1995), the major links of the circuit unquestionably represent a prominent system of connections in the mammalian brain. Hence, the enduring nature of 'Papez's circuit'. Unlike, however, its persistence as anatomical entity, the proposed functional role for the circuit has been less resilient. The early notion that Papez's circuit subserves emotional experience/expression has been abandoned (LeDoux, 1993) and replaced by the proposal that it is primarily involved in mnemonic functions (Aggleton and Brown, 1999). Lesions of each of the major components of the circuit have been shown to disrupt memory (Aggleton and Brown, 1999; Sutherland et al., 1988; Sziklas and Petrides, 1993). The mammillary bodies represent a major output from the hippocampus in Papez's circuit (Amaral and Witter, 1995). It has recently been shown that cells of mammillary body fire rhythmically in bursts synchronous with the theta rhythm of the hippocampus (Bland et al., 1995; Kirk et al., 1996; Kocsis and Vertes, 1994, 1997) and that this rhythmical activity is dependent upon the action of the hippocampus on the mammillary bodies (Bland et al., 1995; Kirk et al., 1996). It is well established that the mammillary bodies project massively to the anterior thalamus (Shibata, 1992), which taken together with the demonstration that mammillary body cells fire synchronously with theta, suggests that the mammillary bodies may act on the anterior thalamus, possibly in the manner that the hippocampus acts on the mammillary bodies, to rhythmically activate cells of the anterior thalamus at theta frequency. We demonstrated that approximately 75% of cells of the anterior ventral nucleus of the thalamus fire rhythmically synchronous with the hippocampal theta rhythm and the activity of 46% of these anterior ventral neurons was highly correlated with theta. These findings, together with demonstration of theta-rhythmically firing cells in other structures of Papez's circuit, indicate that a theta-rhythmic signal may resonate throughout Papez's circuit, possibly involved in the control of mnemonic functions of the circuit.

Stimulus-dependent neuronal oscillations and local synchronization in striate cortex of the alert cat.

Neuronal responses to visual stimuli that are correlated on a millisecond time scale are well documented in several areas of the mammalian visual cortex. This coherent activity often takes the form of synchronous rhythmic discharges ranging in frequency from 20 to 70 Hz. We performed experiments to determine the incidence and properties of this rhythmic activity in the striate cortex of alert cats and to compare this activity to similar data collected in the striate cortex of anesthetized cats. The results demonstrate that optimal visual stimuli evoke robust, locally synchronous, 20-70 Hz oscillatory responses in the striate cortex of cats that are fully alert and performing a visual fixation task. The oscillatory activity is stimulus dependent, largely absent during periods of spontaneous activity, and shows a systematic increase in frequency with increasing stimulus velocity. Thus, the synchronous oscillatory activity observed in this and earlier studies cannot be explained as an artifact of anesthesia nor as a phenomenon that occurs independent of visual stimulation. Rather, it is a robust process that is present in the alert state and is dependent on the presence and specific properties of visual stimuli.

Trigeminal inputs to reticulospinal neurones in lampreys are mediated by excitatory and inhibitory amino acids.

Reticulospinal (RS) neurones integrate sensory inputs from several modalities to generate appropriate motor commands for maintaining body orientation and initiation of locomotion in lampreys. As in other vertebrates, trigeminal afferents convey sensory inputs from the head region. The in vitro brainstem/spinal cord preparation of the lamprey was used for characterizing trigeminal inputs to RS neurones as well as the transmitter systems involved. The trigeminal nerve on each side was electrically stimulated and synaptic responses, which consisted of mixed excitation and inhibition, were recorded intracellularly in the middle and posterior rhombencephalic reticular nuclei. The EPSPs were mediated by activation of alpha-amino-3-hydroxy-5-methyl-4-isoxazolepropionic acid (AMPA)/kainate receptors. An increase in the late phase of the excitatory response occurred when Mg2+ ions were removed from the Ringer's solution. This effect was antagonized by 2-amino-5-phosphonopentanoate (2-AP5) or reversed by restoring Mg2+ ions to the perfusate suggesting the activation of N-methyl-D-aspartate (NMDA) receptors. IPSPs were mediated by glycine. These findings are similar to those reported for other types of sensory inputs conveyed to RS neurones, where excitatory and inhibitory amino acid transmission is also involved.

Relation of olfactory EEG to behavior: time series analysis.

Oscillatory electroencephalographic bursts were measured from 64 electrodes implanted on the olfactory bulbs of rabbits. Oscillatory bursts that occurred before and during presentation of odorant conditioned stimuli (CSs) were selected in brief segments. Comparisons between the 64 traces and their spectra showed that, despite amplitude differences between channels, every burst had a common waveform over the entire array. The spectra showed 2 to 5 distinct peaks in each burst. Each trace was fitted with the sum of 5 cosines to express the burst in ten 8 X 8 matrices of amplitude and phase values at its peak frequencies. Two types of burst were identified. Those with dominant frequencies greater than 55 Hz had one narrow dominant spectral peak and reproducible spatial patterns of its amplitude within subgroups of bursts relating to control and odorant CS conditions. Those with dominant frequencies less than 55 Hz were disorderly; their spectra were broad, and their spatial patterns of amplitude did not reproduce within subgroups. A behavioral assay showed that the high- and not the low-frequency bursts contained odor-specific information.

Conditioning of relative frequency of sniffing by rabbits to odors.

The sniff was identified by a brief episode of increased respiratory rate, usually with a well-defined time of onset. It was detected against the background of respiratory activity in rabbits simply, reliably, and noninvasively by statistical evaluation of digitized pneumograph records. The basal rate of exploratory sniffing was controlled by familiarization. Upon conditioning to olfactory cues, the rate of sniffing for CS+ increased sharply above the basal rate during the first 10 trials and was maintained at high levels by continued reinforcement. During extinction with discrimination between olfactory cues, the rate for CS- fell sharply at first and then more slowly toward the basal rate. With pseudoconditioning, the rabbits responded to an unpaired odor after several sessions; the rates of response acquisition and extinction and the maintained level of responding were lower than with a paired odor in classical delayed conditioning, and the response was not discriminative in respect to another novel odor given during extinction. The sniff displayed a prominent sensory bias for olfactory cues. The relative frequencies of sniffing and respiratory slowing were measured as conditioned responses by screening procedures with a small computer.