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M Penttonen

Publications and source records attributed to M Penttonen.

29 records · Page 2Linked to original sources

Intracellular correlates of hippocampal theta rhythm in identified pyramidal cells, granule cells, and basket cells.

The cellular-synaptic generation of rhythmic slow activity (RSA or theta) in the hippocampus has been investigated by intracellular recording from principal cells and basket cells in anesthetized rats. In addition, the voltage-, coherence-, and phase versus depth profiles were examined by simultaneously recording field activity at 16 sites in the intact rat, during urethane anesthesia, and after bilateral entorhinal cortex lesion. In the extracellular experiments the large peak of theta at the hippocampal fissure was attenuated by urethane anesthesia and abolished by entorhinal cortex lesion. The phase versus depth profiles were similar during urethane anesthesia and following entorhinal cortex lesion but distinctly different in the intact, awake rat. These observations suggest that dendritic currents underlying theta in the awake rat may not be revealed under urethane anesthesia. The frequency of theta-related membrane potential oscillation was voltage-independent in pyramidal neurons, granule cells, and basket cells. On the other hand, the phase and amplitude of intracellular theta were voltage-dependent in all three cell types with an almost complete phase reversal at chloride equilibrium potential in pyramidal cells and basket cells. At strong depolarization levels (less than 30 mV) pyramidal cells emitted calcium spike oscillations, phase-locked to theta. Basket cells possessed the most regular membrane oscillations of the three cell types. All neurons of this study were verified by intracellular injection of biocytin. The observations provide direct evidence that theta-related rhythmic hyper-polarization of principal cells is brought about by the rhythmically discharging basket neurons.(ABSTRACT TRUNCATED AT 250 WORDS)

Anesthesia↗

Hippocampal event-related potentials to pitch deviances in an auditory oddball situation in the cat: experiment I.

Hippocampal event-related potentials (ERP) in the areas CA1, CA3, and dentate fascia (Df) were recorded in cats during an oddball situation when pitch deviant tones occurred in a series of standard tones. When difference waves were calculated by subtracting ERPs to the standard tones from those to the deviant tones, no clear N40d, corresponding to a cat analogue of the human mismatch negativity (MMN) observed in earlier studies, could be detected. Instead, a prominent later negativity (N130d) was observed. A possible extra-hippocampal source of the process reflected by the MMN-like negativity, and a relation between an orienting response (OR) and the N130d are discussed.

Acoustic Stimulation↗

Behavioral and hippocampal evoked responses in an auditory oddball situation when an unconditioned stimulus is paired with deviant tones in the cat: experiment II.

Event-related potentials (ERP) in the areas CA1, CA3 and dentate fascia (Df) of the hippocampal formation were recorded during an oddball situation in the cat. A rewarding electrical stimulation of the lateral hypothalamus (US) was paired with deviant tones (2500 Hz) that occurred randomly in a series of the standard tones (2000 Hz) given to the left ear. In addition to developing orienting head movements to the side of the deviant tones, an increase in the amplitude of parallel hippocampal ERPs was observed. Both the behavioral and neural responses appeared not until a 50 ms latency range. Furthermore, time-amplitude characteristics of the ERPs corresponded to time-acceleration characteristics of the conditioned orienting head movements. The results are discussed in the context of a cat analogue of the human mismatch negativity (MMN) and a role of the hippocampal formation to model and predict the conditioned behavioral orienting responses.

Acoustic Stimulation↗

Hippocampal CA1 interneurons: an in vivo intracellular labeling study.

Fast spiking interneurons in the CA1 area of the dorsal hippocampus were recorded from and filled with biocytin in anesthetized rats. The full extent of their dendrites and axonal arborizations as well as their calcium binding protein content were examined. Based on the spatial extent of axon collaterals, local circuit cells (basket and O-LM neurons) and long-range cells (bistratified, trilaminar, and backprojection neurons) could be distinguished. Basket cells were immunoreactive for parvalbumin and their axon collaterals were confined to the pyramidal layer. A single basket cell contacted more than 1500 pyramidal neurons and 60 other parvalbumin-positive interneurons. Commissural stimulation directly discharged basket cells, followed by an early and late IPSPs, indicating interneuronal inhibition of basket cells. The dendrites of another local circuit neuron (O-LM) were confined to stratum oriens and it had a small but high-density axonal terminal field in stratum lacunosum-moleculare. The fastest firing cell of all interneurons was a calbindin-immunoreactive bistratified neuron with axonal targets in stratum oriens and radiatum. Two neurons with their cell bodies in the alveus innervated the CA3 region (backprojection cells), in addition to rich axon collaterals in the CA1 region. The trilaminar interneuron had axon collaterals in strata radiatum, oriens and pyramidale with its dendrites confined to stratum oriens. Commissural stimulation evoked an early EPSP-IPSP-late depolarizing potential sequence in this cell. All interneurons formed symmetric synapses with their targets at the electron microscopic level. These findings indicate that interneurons with distinct axonal targets have differential functions in shaping the physiological patterns of the CA1 network.

Animals↗

Inhibitory CA1-CA3-hilar region feedback in the hippocampus.

The organization of the hippocampus is generally thought of as a series of cell groups that form a unidirectionally excited chain, regulated by localized inhibitory circuits. With the use of in vivo intracellular labeling, histochemical, and extracellular tracing methods, a longitudinally widespread, inhibitory feedback in rat brain from the CA1 area to the CA3 and hilar regions was observed. This long-range, cross-regional inhibition may allow precise synchronization of population activity by timing the occurrence of action potentials in the principal cells and may contribute to the coordinated induction of synaptic plasticity in distributed networks.

Animals↗

Effects of lateralized US and CS presentations on conditioned head turning and bilateral cingulate cortex responses in cats.

A lateralized tone conditioned stimulus (CS+) paired with a rewarding medial forebrain bundle (MFB) stimulation unconditioned stimulus (US) was presented to one ear of the cat, while the same tone was presented to the other ear alone (CS-). Specifically, the electrical stimulation of the MFB elicited a contralateral head turn as the unconditioned response (UR). Correspondingly, the CS+ was given to an ear contralateral to the direction of the UR. During the CS test session the cats oriented toward the tones, but these head movements habituated rapidly. During conditioning the cats developed stereotypical extended vigorous head turns to the CS+, with significantly greater acceleration and shorter onset latencies to the CS+ than to the CS-. Head turns in response to the CS+ were always ipsilateral to the tone, but responses to the CS- were in some cats ipsilateral and in most cats contralateral to the tone. Recordings of the slow potential responses showed a broad negative deflection in the cingulate cortex, with peaks at 140 and 250 ms. The amplitude of this negative potential to the CS+ was larger than that to the CS-, but no asymmetries were found between the hemispheres. The present behavioral paradigm is potentially useful for studying the neural basis of conditioned approach responses.

Acoustic Stimulation↗

Bilaterally recorded multiple-unit activity of the cingulate cortex during head turning conditioning with unilateral medial forebrain bundle stimulation.

Cats were conditioned to turn their heads using a tone conditioned stimulus (CS) and medial forebrain bundle stimulation (MFB) unconditioned stimulus (US). The CS+ was delivered to one ear at a time, in random order, followed by the US. A tone of a different frequency was used as a CS-. The cats learned to respond differentially to the CSs showing head movements of greater acceleration to the CS+ than CS- over sessions. Bilateral recordings of cingulate cortex multiple-unit activity showed increased response amplitudes over sessions and larger responses in the hemisphere ipsilateral to the US. Since ipsilateral multiple-unit responses did not differ for the CSs, the asymmetry was probably due to the sensitizing effect of the unilateral US. Although increases in cingulate cortex neural activity coincided with increases in conditioned head movements, larger activation of the cingulate cortex ipsilateral to the US suggests that the neural changes were independent of these movements.

Animals↗

Asymmetries in classically conditioned head movements and cingulate cortex slow potentials in cats.

Presuming that conditioned head movements in the cat indicate a preference for a specific direction, asymmetries were also expected to be found in bilaterally recorded cingulate cortex slow potentials to a symmetrical tone stimulus during discriminative conditioning. Conditioned stimuli (CS, 1000 and 2500 Hz tones) were delivered through two miniature loudspeakers fitted to the head in front of both ears. The unconditioned stimulus (US) was an electrical stimulation of either the left or right lateral hypothalamus yielding, in addition to its rewarding effects, an unconditioned head turning response (UR) either to the left, right, forwards or upwards. Discrimination learning appeared as more extended and more rapid headturns and greater negativity in the cingulate cortex to the CS+ than to the CS-. The preferred conditioned behavioral response (CR) was a headturn to the left in 11 of 12 cats accompanying increased negativity in evoked slow potentials in the right cingulate cortex during the 128-324 ms period after the CS onset. No differences in the movements and in the cingulate potentials were found between the left versus right US stimulations. The results showed that regardless of the direction of the original unconditioned head movement, the direction of the CR was at the end of the training period to the left in 11 out of the 12 cats. Larger evoked slow potentials observed in the right cingulate cortex which were not present in the baseline measurements suggest a population bias between the hemispheres.

Animals↗

Conditioned orienting (alpha) and delayed behavioral and evoked neural responses during classical conditioning.

A differentiation of short-latency (alpha) and long-latency (delayed) classically conditioned behavioral and evoked neural (hippocampal) responses was attempted. Further, facilitation and retardation of these responses were studied in an experimental design in which 10 paired conditioning sessions either preceded (CC-CO group) or followed (CO-CC group) 10 randomly unpaired presentations of conditioned stimuli (CS) and unconditioned stimuli (UCS). A 2024-ms tone (1000 Hz) was delivered directly through a miniature earphone to the left ear, eliciting an orienting head movement ('alpha' response) to the left. The unconditioned stimulus (UCS) was a direct 1024-ms stimulation of the lateral hypothalamic area overlapping the CS (delayed paradigm) so that both stimuli terminated simultaneously. The UCS elicited approach behavior and a specific head movement in each animal. The latency and the direction of the head movement were used as criteria for a differentiation of the short-latency and long-latency conditioned responses (CR). All cats showed conditioned short-latency responses. Pairing specific long-latency head movements were observed in 10 of 13 cats and 6 of them showed a long-latency CR which was a head movement to the right, while the short-latency CR on the same trials was a head movement to the left. Hippocampal (subiculum, dentate fascia and CA1) evoked responses also showed pairing specific CRs appearing as increased negativity (short-latency CR), or increased positivity (long-latency CR). Additional reversed stimulus order (backward) sessions supported an assumption of the different nature of the short-latency and long-latency CRs: the long-latency CRs showed extinction while the short-latency CRs remained. The unpaired pre-exposure to the CSs and UCSs in the CO-CC group resulted in the retarded acquisition of the behavioral responses during the subsequent paired sessions.

Animals↗

Behavioral and neural characteristics of short-latency and long-latency conditioned responses in cats.

Head movements to the conditioned stimulus (a tone CS to the left ear) were studied in 6 cats. An attempt was made to differentiate an orienting, short-latency (alpha) response from the long-latency conditioned (delayed) response. The unconditioned stimulus (UCS) was a brain stimulation to the lateral hypothalamus eliciting, in addition to orienting and approach behavior, a specific, stereotypical head movement. These behavioral characteristics of the unconditioned head movement were used for differentiating it from the conditioned short-latency head movement. Paired conditioning and randomly unpaired control sessions (5 daily sessions each) were given in balanced order to each animal. Evoked neural responses in the hippocampus and cingulate cortex were recorded simultaneously to compare the time-amplitude characteristics of evoked responses to earlier findings in multiple-unit recordings. The results supported the differentiation of the behavioral responses. The time-amplitude course of the evoked neural responses showed complex changes, appearing as an increase in the negativity during the alpha-response period (150- to 450-ms interstimulus interval [ISI]) and as an increase in the positivity during the long-latency period (700- to 1,000-ms ISI + 150- to 450-ms UCS period) on omitted-UCS (CS-alone test) trials.

Alpha Rhythm↗