Pleasure: The missing link in the regulation of sleep.

Although largely unrecognized by sleep scholars, sleeping is a pleasure. This report aims first, to fill the gap: sleep, like food, water and sex, is a primary reinforcer. The levels of extracellular mesolimbic dopamine show circadian oscillations and mark the "wanting" for pro-homeostatic stimuli. Further, the dopamine levels decrease during waking and are replenished during sleep, in opposition to sleep propensity. The wanting of sleep, therefore, may explain the homeostatic and circadian regulation of sleep. Accordingly, sleep onset occurs when the displeasure of excessive waking is maximal, coinciding with the minimal levels of mesolimbic dopamine. Reciprocally, sleep ends after having replenished the limbic dopamine levels. Given the direct relation between waking and mesolimbic dopamine, sleep must serve primarily to gain an efficient waking. Pleasant sleep (i.e. emotional sleep), can only exist in animals capable of feeling emotions. Therefore, although sleep-like states have been described in invertebrates and primitive vertebrates, the association sleep-pleasure clearly marks a difference between the sleep of homeothermic vertebrates and cool blooded animals.

Effects of the muscarinic agonist pilocarpine on vigilance states and locomotor activity in ring doves.

Cholinergic systems play a significant role in regulating a variety of behavioral functions in mammals and birds. The aim of this work is to study the effects of the muscarinic agonist pilocarpine on behavioral states by visual inspection and electroencephalographic recording; also, locomotor activity was continuously recorded by infrared interruption system in ring doves. The current results in birds demonstrated that the muscarinic agonist pilocarpine (1 and 3mg/kg, i.p.) primarily induced theta activity in addition to promote passive waking, while diminished active waking, the EEG slow wave rhythm and REM sleep in ring doves. The locomotor activity recorded continuously in ring doves diminished after pilocarpine treatment, which was in good agreement with the observed reduction of active waking derived of the EEG study. Altogether, the current results are similar to the effects of pilocarpine previously reported in mammals. In conclusion, hippocampal theta rhythm in birds suggests that this rhythm is an ancestral property of hippocampal function and similar cholinergic mechanisms regulate vigilance states and theta generation in mammals and birds.

Comments on evolution of sleep and the palliopallial connectivity in mammals and birds.

This commentary is referred to the review signed by Rattemborg [N.C. Rattenborg, Evolution of slow wave sleep and palliopallial connectivity in mammals and birds. A hypothesis. Brain Res. Bull. 69 (2006) 20-29]. We propose that the review missed important aspects in relation to the characteristics of sleep in poikilotherm vertebrates and in the evolution of sleep. Poikilotherms continuously show an EEG dominated by slow waves, but its highest amplitude appears not during sleep, but during active waking. In addition, they show an arousal reaction which consists in an increase in EEG amplitude and synchrony, opposite to mammals and birds. As a consequence, most of the conclusions proposed in the review should be rejected.

[The work of Santiago Ramón y Cajal in the Revista Trimestral Micrográfica (Trabajos del Laboratorio de Investigaciones Biológicas)]. / La obra de Santiago Ramón y Cajal en Revista Trimestral Micrográfica (Trabajos del Laboratorio de Investigaciones Biológicas).

AIM: This paper is based on a study of Revista Trimestral Micrografica (Trabajos del Laboratorio de Investigaciones Biologicas) between its creation by Santiago Ramon y Cajal in 1896 and his death in 1934. DEVELOPMENT: The journal Revista Trimestral Micrografica was the main way in which Santiago Ramon y Cajal and his school published their work since its creation. Ramon y Cajal created the journal for two main reasons: first, he needed a rapid system to publish his own work; second, the journal could serve to encourage his pupils. The journal published many important reports defending the neuronal theory which expanded the cellular one to include the nervous system.

Dualism and uniformism in sleep.

The phenomenological evidence for distinguishing between REM and NREM sleep is overwhelming. However, this difference has only been found thanks to electrophysiological analytical methods, and is practically non existent in phenotypic terms, i.e., observable with the naked eye. It is well accepted that the selective pressure determining evolutionary changes can only work upon phenotypic differences. Hence, it follows that the differences between REM and NREM could not have been selected through evolution and this implies that, in functional terms, both states could be equivalent.

Why we sleep: the evolutionary pathway to the mammalian sleep.

The cause of sleep is a complex question, which needs first, a clear distinction amongst the different meanings of a causal relationship in the study of a given behavior, second, the requisites to be met by a suggested cause, and third, a precise definition of sleep to distinguish behavioral from polygraphic sleep. This review aims at clarifying the meaning of the question and at showing the phylogenetic origin of the mammalian and avian sleep. The phylogenetic appearance of sleep can be approached through a study of the evolution of the vertebrate brain. This began as an undifferentiated dorsal nerve, which was followed by the development of an anterior simplified brain and ended with the formation of the multilayered mammalian neocortex or the avian neostriate. The successive stages in the differentiation of the vertebrate brain produced, at least, two different waking types. The oldest one is the diurnal activity, bound to the light phase of the circadian cycle. Poikilotherms control the waking from the whole brainstem, where their main sensorymotor areas lie. Mammals developed the thalamocortical lines, which displaced the waking up to the cortex after acquiring homeothermy and nocturnal lifestyle. In order to avoid competence between duplicate systems, the early waking type, controlled from the brainstem, was suppressed, and by necessity was turned into inactivity, probably slow wave sleep. On the other hand, the nocturnal rest of poikilotherms most probably resulted in rapid eye movement sleep. The complex structure of the mammalian sleep should thus be considered an evolutionary remnant; the true acquisition of mammals is the cortical waking and not the sleep.

Nonlinear, fractal, and spectral analysis of the EEG of lizard, Gallotia galloti.

Electroencephalogram (EEG) from dorsal cortex of lizard Gallotia galloti was analyzed at different temperatures to test the presence of fractal or nonlinear structure during open (OE) and closed eyes (CE), with the aim of comparing these results with those reported for human slow-wave sleep (SWS). Two nonlinear parameters characterizing EEG complexity [correlation dimension (D2)] and predictability [largest Lyapunov exponent (lambda(1))] were calculated, and EEG spectrum and fractal exponent beta were determined via coarse graining spectral analysis. At 25 degrees C, evidence of nonlinear structure was obtained by the surrogate data test, with EEG phase space structure suggesting the presence of deterministic chaos (D2 approximately 6, lambda(1) approximately 1. 5). Both nonlinear parameters were greater in OE than in CE and for the right hemisphere in both situations. At 35 degrees C the evidence of nonlinearity was not conclusive and differences between states disappeared, whereas interhemispheric differences remained for lambda(1). Harmonic power always increased with temperature within the band 8-30 Hz, but only with OE within the band 0.3-7.5 Hz. Qualitative similarities found between lizard and human SWS EEG support the hypothesis that reptilian waking could evolve into mammalian SWS.

Interhemispheric differences in awake and sleep human EEG: a comparison between non-linear and spectral measures.

Interhemispheric differences in the EEG of nine healthy right-handed human subjects (C3 vs. C4 derivations) were investigated during resting wake with closed eyes (CE) and sleep stages I, II, III, IV and REM. The harmonic power spectral density within the EEG main spectral bands, the fractal (Dr) and the correlation (D2) dimension as well as the largest Lyapunov exponent (lambda1) of both hemispheres were compared. In addition, the relationships between non-linear and spectral measures were analyzed. Dr, D2, lambda1 and the power in alpha band exhibited interhemispheric differences during waking, the values from the right hemisphere (RH) being higher than those of the left (LH) except for lambda1. During slow wave sleep (SWS), non-linear parameters detected opposite EEG asymmetries (D2 in stage III and lambda1 in stage IV) to those found in the other behavioural stages. In addition, both D2 and lambda1 were correlated (negatively) with the power in the delta band, but lambda1 was also correlated (positively) with the power in the alpha and beta bands. In conclusion, RH appears to be more complex though more predictable than the LH during CE and sleep stages I and II, these characteristics changing to the LH during SWS.

Thin-layer chromatographic determination of catecholamines, 5-hydroxytryptamine, and their metabolites in biological samples: a review.

A review of methodology for separation, detection, and quantitative determination of catecholamines, 5-hydroxytryptamine, and their acidic metabolites in biological tissue and fluids by thin-layer chromatography is presented. Selected procedures, including fluorometric scanning densitometry for catecholamine acetyl derivatives and color scanning densitometry for acids, are described.

Main trends in rectal temperature during sleep. Hypnogram and thermogram.

Changes in sleep state and night-time body temperature were studied in 13 healthy young men. Temperature data showed serial dependence that was removed when considering three main segments of the thermogram: an initial rapid drop between 60 and 140 min after lights off, followed by a slower decrease to the thermal minimum and a final temperature increase until waking. The first segment was fitted by a two-term function, while the other two segments were fitted by linear regression. Temperature data obtained after fitting these models were stationary and used to calculate the fast Fourier transformation and the cross-correlation functions between the hypnogram and the thermogram of each subject. Attaining stationarity has unmasked short oscillations during sleep and some temperature series showed the presence of ultradian oscillations with a period of 90-100 min. Significant cross-correlations between sleep stages and body temperature were found, and the parameters of the two-term function fitted to the first temperature drop were related to the sign of the correlation and to that time series (sleep or temperature) which preceded the other during the night.

A simple thin-layer chromatographic method for the analysis of buprenorphine in urine.

A simple and sensitive (at the ng/mL level) method for buprenorphine assay in urine is described. Buprenorphine is extracted by a C18 cartridge, derivatized with dansyl chloride, resolved by a uni-dimensional, two developing mobile phase high-performance thin layer chromatography system and quantified by fluorescence densitometry at 340 nm.

Thin-layer chromatographic determination of brain catecholamines and 5-hydroxytryptamine.

A simple and sensitive (up to nanogram level) method to determine norepinephrine, serotonin and dopamine in rat brain is described. The amines are acetylated and the derivatives are resolved by thin layer chromatography (TLC) on silica high performance thin layer chromatography (HPLC) plates. Quantification is achieved by fluorescence densitometry at 415 nm excitation wavelength.

A simple method for plasma cannabinoid separation and quantification.

A sensitive (up to nanogram level) method for resolving a cannabinoid mixture in plasma is described. Cannabinoids were extracted with a C-18 Sep Pak cartridge and derivatized with dansylchloride. Then the derivatives were resolved on thin layer HPTLC silica plates which were developed and quantified by fluorescence densitometry at 340 nm.

Histaminergic drugs in the rat caudate nucleus: effects on learned helplessness.

The effect of pharmacological manipulation of histaminergic receptors in the caudate nucleus (CN) has been examined in rats previously submitted to inescapable electric shock to produce learned helplessness (LH). Histamine H1 agonist 2-tiazolylethyl amine (TEA) microinjection produced protective effects, preventing the activity and cognitive loss typical in LH. Injection of the H1 antagonist astemizole (AZ) produced effects symmetrical to those produced by TEA, further reducing activity and impairing cognitive functions. The histamine H2 agonist 4-methyl-histamine (4MH) produced a shift on the side preference for rotation that interfered in the learning tests and obscured the effects of this drug on LH. Injection of the H2 antagonist cymetidine (CYM) caused LH-like effects in control animals. Thus, brain histamine seems to play a relevant role in the control of motor and cognitive functions of the CN.

On the evolution of waking and sleeping.

1. The aim of this paper is to present a new hypothesis to explain the evolution of the sleeping and waking states. 2. We propose that the reptilian waking state and the mammalian slow wave sleep are homologous states. 3. We also propose that instead of looking at the polygraphic sleep as a new evolutive acquisition of mammals and birds, it seems more convenient to look at the full waking state; the "advanced wakefulness" as the true new evolutionary acquisition of these animals. 4. These conclusions are reached after examining some available reports of slow wave electroencephalogram in waking reptiles, some other reports showing signs of rapid eye movement sleep in this same group and the coevolution between sleep states and thermoregulation. Finally, a clear parallelism between sleep ontogeny and phylogeny is shown under the light of the proposed hypothesis.

Morphofunctional changes in gastrointestinal tract of rats due to cafeteria diet.

Female rats fed a cafeteria diet from birth developed obesity at 60 days of age and their stomach, small intestine and caecum were enlarged when compared with controls, i.e. these regions had greater food storage capacity. In spite of the enlargement, these regions had similar or reduced weight and linear density, which is seen as proof of reduced mechanical performances. Cafeteria diet produced increased glucose duodenal absorption in older animals unlike the typical reduction known in controls. Tryptophan absorption was maintained high in adulthood, compensating for the low structural nutritive properties of the cafeteria diet. The results are interpreted as an adaptation to the cafeteria diet effects and properties: the characteristic overeating of foodstuffs with greater energy density, lower mechanical requirements and lower structural nutritive value than pelleted chow.