Anterograde tracers, Phaseolus vulgaris leucoagglutinin (PHA-L) and horseradish peroxidase (HRP), were used to study the thalamocortical afferents of the posteromedial barrel subfield (PMBSF) in rat primary somatosensory cortex (SI) at both light- and electron-microscopic levels. The PMBSF, also known as the barrel cortex, can be subdivided into barrel and interbarrel areas on the basis of cytoarchitectonic characteristics. Restricted injections confined to either the ventroposterior medial (VPM) or the rostral part of the posterior (Pom) nucleus allowed us to study and compare their projection patterns to the barrel cortex. We found that the interbarrel area receives inputs exclusively from the Pom, whereas the barrel area receives inputs from both the Pom and VPM. The laminar distributions of these two projections are largely segregated. After an injection of PHA-L or HRP into the VPM, labeled bouton-like swellings are found in layer VI and in layers IV through I of the barrel area, with the highest concentration in layer IV. On the other hand, after an injection of PHA-L or HRP into the Pom, labeled bouton-like swellings are distributed from upper layer V to layer I of the interbarrel area, as well as in layers V and I of the barrel area. Ultrastructural analysis showed that labeled bouton-like swellings of the VPM and the Pom pathways make synaptic contacts onto cortical neurons, and that these contacts are asymmetrical. Therefore, the VPM and the Pom projections are complementary to each other in the barrel cortex, and together they provide thalamic inputs to most layers of both the barrel and interbarrel areas. The differential patterns of terminations of the VPM and the Pom projections in the barrel cortex suggest that they may be involved in different types of cortical processing. Furthermore, our present findings may provide the anatomical basis for two parallel thalamocortical pathways, which previous physiological studies have indicated are each concerned with particular submodalities of somatic information.
SUMMARY AND CONCLUSIONSI. Thalamic relay cells, including those of the lateral geniculate nucleus, display a low-threshold spike (LT spike), which is a large depolarization due to an increased Ca2+ conductance. Typically riding the crest of each LT spike is a burst of from two to seven action potentials, which we refer to as the LT burst. The LT spike is voltage dependent, because if the cell's resting membrane potential is more depolarized than roughly -60 mV, the LT spike is inactivated, but if more hyperpolarized, the spike is deinactivated and can be activated by a depolarization, such as from an afferent excitatory postsynaptic potential (EPSP). Thalamic relay cells thus display two response modes: a relay or tonic mode, when the cell is depolarized and LT spikes are inactivated, leading to tonic firing of action potentials; and a burst mode, when the cell is hyperpolarized and tends to respond with LT spikes and their associated bursts of action potentials.2. We were interested in the contribution of the LT spike on the transmission of visually evoked signals through geniculate relay cells to visual cortex. We recorded intracellularly from geniculate cells in an anesthetized, paralyzed, in vivo cat preparation to study the effects of membrane voltage, and thus the presence or absence of LT spikes, on responses to drifting sine-wave gratings. We monitored the visually evoked responses of 14 geniculate neurons (6 X, 7 Y, and 1 unclassified) at different membrane potentials at which LT spikes were inactivated or deinactivated.3. Changing membrane voltage during visual stimulation switched the response mode of every cell between the relay and burst modes. In the burst mode, LT spikes occurred in phase with the visual stimulus and not at rhythmic intervals uncorrelated to visual stimuli. To any given stimulus cycle, the cell responded usually with an LT burst or a tonic response, and rarely was more than one LT burst evoked by a stimulus cycle. Occasionally a single cycle evoked both an LT burst and tonic response, but always the LT burst occurred first.4. The spatial tuning characteristics of the cells did not differ dramatically as a function of membrane potential, because the tuning of the LT bursts was quite similar to that of the tonic response component. Although we did not obtain complete temporal tuning properties, we did note that hyperpolarized cells responded reliably with LT bursts at several temporal frequencies.5. A consistent difference was seen between the LT burst and tonic response components in terms of response linearity. We measured this by computing the fundamental and second harmonic Fourier amplitudes of the responses (Fl and F2, respectively ). The Fl amplitude represents the linear portion of the response, and the F2 amplitude represents a measure of response nonlinearity. We found that, for every cell, the F2-to-F1 ratio was considerably higher for the LT burst than for the tonic response component.6. We found that the bursts associated with LT spikes had interspike intervals 54 ms. H...
1. In an anesthetized, paralyzed in vivo preparation, we recorded extracellular responses of 61 geniculate neurons (2 W, 25 X, 33 Y, and 1 mixed) to drifting sine-wave gratings of various spatial frequency, temporal frequency, and contrast. Our goal was to study the differential contributions to these visual responses of bursting caused by voltage dependent, low-threshold (LT) Ca2+ spikes and of purely tonic responses unrelated to LT spikes. Cells responding with LT spikes are said to be in the burst firing mode and those responding in a purely tonic fashion to be in the relay or tonic firing mode. We separated the total visual response into LT burst and tonic components by use of the empirical criteria set forth in our intracellular study described in the previous paper (Lu et al. 1992). A response component was considered to be an LT burst if its action potentials displayed interspike intervals < or = 4 ms and if the first spike in the burst episode occurred after a silent period of > or = 100 ms (or > or = 50 ms when the neuron responds to visual stimuli at temporal rates > or = 8 Hz). All other activity is considered to be part of the tonic response. 2. In addition to LT bursts, we recognized another type of burst response, the high-threshold (HT) burst. These also have clusters of action potentials with interspike intervals < or = 4 ms. However, HT bursts, unlike LT bursts, lack a preburst silent period. HT bursts are part of the tonic response component and merely reflect the gradual decrease in interspike intervals that occurs as the cell becomes more depolarized and thus more responsive. Thus interspike interval is a necessary but insufficient criterion to identify LT bursts. 3. Visually evoked LT bursts were recorded among W, X, and Y cells. When evoked, LT bursts occurred in phase with drifting sine-wave grating stimuli at a rate never exceeding one per stimulus cycle. In response to individual cycles of the visual stimulus, LT bursts could comprise the total response, a tonic component could comprise the total response, or an LT burst and tonic component could be mixed. When a stimulus evoked a mixture of LT bursts and tonic response components, LT bursts were always the first response. 4. Of the 61 cells tested with grating stimuli, 47 exhibited LT bursts and 14 did not. Those that did exhibited varying amounts of burstiness.(ABSTRACT TRUNCATED AT 400 WORDS)
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