Effects of altered photoperiod on circadian clock and lipid metabolism in rats

Disruption of circadian clock timekeeping due to changes in the photoperiod enhances the risk of lipid metabolism disorders and metabolic syndrome. However, the effects of altered photoperiods on the circadian clock and lipid metabolism are not well understood. To explore the effects of altered photoperiods, we developed a rat model where rats were exposed to either short-day or long-day conditions. Our findings demonstrated that altered photoperiods mediated circadian clocks by partly disrupting rhythmicity and shifting phase values of clock genes. We also showed that compared to long-day conditions, rats under short-day conditions exhibited more photoperiodic changes in a variety of physiological outputs related to lipid metabolism, such as significant increases in serum triglyceride (TG), high-density lipoprotein, and leptin levels, as well as increased body weight, fat:weight ratio, and hepatic TG levels. These increments were gained possibly through upregulated expression of forkhead box O1 (FoxO1), which partly mediates the expression of peroxisome proliferator-activated receptorα (PPARα) to increase the expression of phosphoenolpyruvate carboxykinase (PEPCK), peroxisome proliferator-activated receptor-g coactivator-1β (PGC1β), and fatty acid synthase (Fasn). In addition, the oscillation rhythms of FoxO1, PEPCK, PGC1β, and Fasn expression levels in the livers of rats exposed to a short-day photoperiod were more robust than those exposed to a long-day photoperiod. These findings suggest that a change in photoperiod can partly disrupt the circadian rhythmcity of clock genes, impair lipid metabolism, and promote obesity.     

Effects of prolactin and experimental handling on liver, fat body and ovary lipid contents and their daily variations in Rana esculenta (L.)

Three groups of females of R. esculenta were kept under constant temperature and photoperiod conditions (L:D = 12:12). One group consisted of intact frogs, while the remainder were given saline or prolactin (PRL) injections 6 hours after light onset. After 5 days of such treatment examination was made every 6 hours over a 24-hour period of lipid content in the liver, fat bodies (FBs) and ovaries and the level and composition of plasma lipids. Both the experimental handling and PRL treatment caused in animals a decrease in weight of the liver and mobilization of lipid metabolism, inducing a shift in lipids and a change in their distribution within the organs examined. In animals injected with saline lipids were transported chiefly to FBs, whereas in frogs given PRL the whole, combined lipid content in these organs decreased considerably, suggesting that they had been transported to the tissues and that their catabolism had been intensified. It is suggested also that PRL has sparing effect on the ovarian lipid pool. In both groups, especially in frogs given PRL, there was a marked increase in plasma lipid concentration and the contents in percentages of its different classes. Increase in concentration of phospholipids (3X) and cholesterol (10X) in animals given PRL is particularly distinct. Both experimental handling and PRL administration affected the pattern of diurnal fluctuations in the lipid content of the organs examined.     

Influence of Photoperiod and Leaf Age on Crassulacean Acid Metabolism in Portulacaria afra (L.) Jacq

The possibility that Crassulacean acid metabolism (CAM) is subject to long day photoperiodic control in Portulacaria afra (L.) Jacq., a facultative CAM plant, was studied. Periodic measurements of ¹⁴CO₂ uptake, stomatal resistance, and titratable acidity were made on plants exposed to long and short day photoperiods. Results indicates that waterstressed P. afra had primarily nocturnal CO₂ uptake, daytime stomatal closure, and a large diurnal acid fluctuation in either photoperiod. Mature leaf tissue from nonstressed plants under long days exhibited a moderate diurnal acid fluctuation and midday stomatal closure. Under short days, there was a reduced diurnal acid fluctuation in mature leaf tissue. Young leaf tissue taken from nonstressed plants did not utilize the CAM pathway under either photoperiod as indicated by daytime CO₂ uptake, lack of diurnal acid fluctuation, and incomplete daytime stomatal closure.

The induction of CAM in P. afra appears to be related to the water status of the plant and the age of the leaf tissue. The photosynthetic metabolism of mature leaves may be partly under the control of water stress and of photoperiod, where CAM is favored under long days.

     

Effects of pinealectomy on pituitary gonadotrophs, pituitary gonadotropin potency and hypothalamic gonadotropin releasing activity in Notemigonus crysoleucas

The effects of pinealectomy on pituitary gonadotrophs, pituitary gonadotropin potency and hypothalamic gonadotropin releasing activity were examined in the cyprinid teleost, Notemigonus crysoleucas, exposed to various photoperiod-temperature regimes. In fish exposed to a long photoperiod-warm temperature regime, pinealectomy resulted in a decrease in gonadal activity, in hypothalamic gonadotropin releasing activity and an increase in pituitary gonadotropin potency. Fewer gonadotrophs were present in the pituitary of sham operated fish than in the pituitary of pinealectomized fish. Ovarian development was more rapid in sham operated than in pinealectomized fish exposed to a long photoperiod–low temperature regime. Pituitary gonadotropin activity was also greater in shams than in pinealectomized fish. A short photoperiod-warm temperature regime retarded ovarian development in N. crysoleucas. Pinealectomy reversed this trend. Gonadotrophs made up a greater area of the pituitary in pinealectomized fish than in shams under these conditions. Gonadotropin potency of the pituitary and hypothalamic gonadotropin releasing activity were also greater in pinealectomized fish than in shams. The area of the pituitary occupied by gonadotrophs was greater in pinealectomized than in sham operated animals maintained on a short photoperiod-low temperature regime. Pituitary gonadotropin activity was also greater in pinealectomized fish as compared to shams. Pituitary gonadotropin potency varies diurnally in animals maintained on both short and long photoperiods; the rhythm of variation differs depending on photoperiod. Pinealectomy alters the diurnal rhythm of pituitary gonadotropin potency in animals exposed to both long and short photoperiods. It is concluded that pinealectomy has a pronounced effect on reproductive activity in N. crysoleucas. The effects of pinealectomy on reproduction vary with photoperiod, but are mediated via the hypothalamus and pituitary. In fish exposed to long daylengths the pineal favours reproductive activity, but the epiphysis retards reproductive processes in animals maintained on short photoperiods.     

The effects of pinealectomy on pituitary and plasma gonadotropin levels in Carassius auratus exposed to various photoperiod–temperature regimes

The effects of pinealectomy on pituitary and plasma gonadotropin levels and gonadal development in female goldfish exposed to various photoperiod-temperature regimes during different seasons were examined. Pinealectomy during autumn had no effect on either pituitary or plasma hormone levels or gonadal development. When goldfish are pinealectomized in spring and exposed to long photoperiod conditions, the ovaries regress and plasma gonadotropin levels are significantly depressed compared to sham operated animals. Sham operated goldfish exposed to short photoperiod conditions in spring had regressing ovaries whereas pinealectomized animals under this regime either spawned or had ovaries in the late vitellogenic phase. Plasma gonadotropin titres in the pinealectomized group were significantly lower than those of sham operated animals. The pineal can be either stimulatory or inhibitory to gonadal development depending on the photoperiod regime to which the animals are exposed. The pineal apparently influences gonadal activity by modulating gonadotropin secretion. A diurnal variation in plasma gonadotropin levels was also observed in both sham operated and pinealectomized goldfish exposed to a long photoperiod warm-temperature regime in spring.     

Changes in the lipid composition of Acanthephyra purpurea Milne Edwards (Crustacea: Decapoda) during its diurnal migration: A preliminary investigation

The lipids of the mesopelagic decapod Acanthephyra purpurea Milne Edwards have been studied in relation to its diurnal vertical migration. The total lipid content increased between midnight and early morning, and both the triglyceride and wax ester fractions showed considerable variation in amount. There was a significant increase in the phospholipid content during its period in deep water, suggesting a diurnal rhythm of lipid metabolism associated with the migratory rhythm.     

Suppression of the prolactin surges in the pseudopregnant rat by urethane anesthesia

In pseudopregnancy of the rat prolactin (PRL) is released in the form of twice daily surges (nocturnal and diurnal surges). An attempt was made to examine the effects of urethane anesthesia on PRL surges during pseudopregnancy of the rat. In a preliminary study, using the continuous blood sampling method, the nocturnal PRL surge was completely blocked when urethane (1.0g/kg BW) was administered at 0:00 hr. Urethane (1.0g/kg BW) was injected at 0:00 or 12:00 hr, and serum and pituitary PRL concentrations were measured at 6:00 or 18:00 hr, respectively, to study the effects of urethane on nocturnal or diurnal PRL surges. There were no serum PRL surges during either the nocturnal or diurnal periods following urethane injection. The experiment examining pituitary PRL concentration at 6:00 or 18:00 hr confirmed that urethane (1.0g/kg) anesthesia suppressed the release of PRL surge from the pituitary.     

Morning pulse of prolactin maintains seasonal quiescence in the tammar, Macropus eugenii

The pulse of prolactin, present in female tammars kept in an inhibitory photoperiod and absent 5 days after a change to a stimulatory photoperiod, was artificially maintained during stimulatory photoperiods (Exps 1A and 1B) or abolished during an inhibitory photoperiod (Exp. 2). In Exp. 1A, thyrotrophin-releasing hormone (TRH) was injected i.v. for 8 consecutive mornings and in Exp. 1B ovine prolactin was injected i.v. for 10 consecutive mornings beginning on the day the photoperiod was changed from 15L:9D to 12L:12D. In Exp. 1A, in 5 of 6 control females the prolactin pulse was present until the 3rd morning after the lighting was changed, and absent thereafter. In both experiments the early progesterone peak and the interval to birth in the treated females was significantly delayed compared to the control females; reactivation of the CL did not occur until treatment ceased. In Exp. 2, for 5 consecutive mornings, before the prolactin pulse was expected to occur, bromocriptine was injected i.m. In 2/4 treated animals the prolactin pulse was abolished and these animals showed an early progesterone peak and gave birth 28 and 29 days after treatment began. The remaining 2 animals and the controls (N = 4) did not show reactivation until after they experienced a stimulatory photoperiod. The results indicate that the prolactin pulse is the critical factor in the maintenance of the inhibition of the CL of the tammar during seasonal quiescence.     

Photoperiod affects the diurnal rhythm of hippocampal neuronal morphology of siberian hamsters

Individuals of many species can regulate their physiology, morphology, and behavior in response to annual changes of day length (photoperiod). In mammals, the photoperiodic signal is mediated by a change in the duration of melatonin, leading to alterations in gene expressions, neuronal circuits, and hormonal secretion. The hippocampus is one of the most plastic structures in the adult brain and hippocampal neuronal morphology displays photoperiod-induced differences. Because the hippocampus is important for emotional and cognitive behaviors, photoperiod-driven remodeling of hippocampal neurons is implicated in seasonal differences of affect, including seasonal affective disorder (SAD) in humans. Because neuronal architecture is also affected by the day-night cycle in several brain areas, we hypothesized that hippocampal neuronal morphology would display a diurnal rhythm and that day length would influence that rhythm. In the present study, we examined diurnal and seasonal differences in hippocampal neuronal morphology, as well as mRNA expression of the neurotrophic factors (i.e., brain-derived neurotrophic factor [Bdnf], tropomyosin receptor kinase B [trkB; a receptor for BDNF], and vascular endothelial growth factor [Vegf]) and a circadian clock gene, Bmal1, in the hippocampus of Siberian hamsters. Diurnal rhythms in total length of dendrites, the number of primary dendrites, dendritic complexity, and distance of the furthest intersection from the cell body were observed only in long-day animals; however, diurnal rhythms in the number of branch points and mean length of segments were observed only in short-day animals. Spine density of dendrites displayed diurnal rhythmicity with different peak times between the CA1 and DG subregions and between long and short days. These results indicate that photoperiod affects daily morphological changes of hippocampal neurons and the daily rhythm of spine density, suggesting the possibility that photoperiod-induced adjustments of hippocampal neuronal dynamics might underlie seasonal difference of affective responses. Bmal1 mRNA showed a diurnal rhythm and different expression levels between long and short days were observed. However, there were no strong effects of day length on Bdnf. trkb, and Vegf gene expression, suggesting that these genes are not involved in the photoperiodic effects on hippocampal neurons.     

Seasonal and individual variability of lipid reserves in <Emphasis Type="Italic">Oithona similis</Emphasis> (Cyclopoida) in an Arctic fjord

The lipid storage of the cyclopoid copepod Oithona similis was investigated from spring to late summer 2007 in Kongsfjorden (Svalbard, Norway). The volume of lipid droplets in each individual reflected the amount of stored wax esters. Seasonal changes of lipid storage coupled with informative inter-individual variability were thus obtained. The seasonal pattern showed an increase in lipid store during the spring bloom, starting before the chlorophyll a maximum for both copepodids stage V and females. Those reserves were used during the main reproductive event in June. Individual variability was very high, with a significant proportion of copepods having no droplet, while others were lipid rich. Because of the overlap of generation, females could have different age and feeding history, particularly in September. Consideration of intra-population variability in lipid storage using an optical approach has been shown to be important to understand O. similis’s ecology and life cycle.     

Phase and period responses to short light pulses in a wild diurnal rodent,Funambulus pennanti

Photic phase response curves (PRCs) have been extensively studied in many laboratory-bred diurnal and nocturnal rodents. However, comparatively fewer studies have addressed the effects of photic cues on wild diurnal mammals. Hence, we studied the effects of short durations of light pulses on the circadian systems of the diurnal Indian Palm squirrel, Funambulus pennanti. Adult males entrained to a light–dark cycle (12?h–12?h) were transferred to constant darkness (DD). Free-running animals were exposed to brief light pulses (250 lux) of 15?min, 3 circadian hours (CT) apart (CT 0, 3, 6, 9, 12, 15, 18 and 21). Phase shifts evoked at different phases were plotted against CT and a PRC was constructed. F. pennanti exhibited phase-dependent phase shifts at all the CTs studied, and the PRC obtained was of type 1 at the intensity of light used. Phase advances were evoked during the early subjective day and late subjective night, while phase delays occurred during the late subjective day and early subjective night, with maximum phase delay at CT 15 (?2.04?±?0.23?h), and maximum phase advance at CT 21 (1.88?±?0.31?h). No dead zone was seen at this resolution. The free-running period of the rhythm was concurrently lengthened (deceleration) during the late subjective day and early subjective night, while period shortening (acceleration) occurred during the late subjective night. The maximum deceleration was noticed at CT 15 (?0.40?±?0.09?h) and the maximum acceleration at CT 21 (0.39?±?0.07?h). A significant positive correlation exists between the phase shifts and the period changes (r?=?0.684, p?=?0.001). The shapes of both the PRC and period response curve (τRC) qualitatively resemble each other. This suggests that the palm squirrel’s circadian system is entrained both by phase and period responses to light. Thus, F. pennanti exhibits robust clock-resetting in response to light pulses.     

Endogenous circannual cycles of ovarian activity and changes in prolactin and melatonin secretion in wild and domestic female sheep maintained under a long-day photoperiod

The present study examines the ovulatory activity of wild and domesticated ewes subjected to either a constant photoperiod of long days (16L:8D) or natural changes in daily photoperiod for 16 mo. The aim was to determine whether an endogenous reproductive rhythm controls seasonal reproductive activity in these sheep, and how the photoperiod might affect this. The effects of long-day photoperiods on long-term changes in prolactin and melatonin secretion were also evaluated. The two species showed changes in reproductive activity under the constant photoperiod conditions, suggesting the existence of an endogenous rhythm of reproduction. This rhythm was differently expressed in the two types of ewe (P < 0.05), with the domestic animals exhibiting much greater sensitivity to the effects of long days. A circannual rhythm of plasma prolactin concentration was also seen in both species and under both photoperiod conditions, although in both species the amplitude was always lower in the long-day animals (P < 0.01). The duration of the nocturnal melatonin plasma concentrations reflected the duration of darkness in both species and treatments. The peak melatonin concentration did not differ between seasons either under natural or long-day photoperiods.     

Pituitary prolactin concentrations associated with daily prolactin surges in pseudopregnant rats

During pseudopregnancy (PSP) two surges of prolactin (PRL) secretion from the pituitary are observed, the nocturnal surge at dawn and the diurnal surge in the evening. An attempt was made to clarify the correlation between changes in serum and pituitary PRL concentrations on day 5-6 of PSP. During the nocturnal surge, pituitary PRL concentration decreased significantly from 0000 hr to 0300-0600 hr. On the other hand, the high pituitary PRL concentration remained unchanged during the diurnal surge from 1200 hr to 1800 hr. These findings suggest that the nocturnal and diurnal PRL surges are regulated by separate controlling mechanisms.     

Alterations in the lipid content of pituitary gland and serum prolactin and growth hormone in cadmium treated rats

The present study was undertaken to assess whether chronic exposition to cadmium (Cd, 0.133 mM per liter for 2 months) through drinking water may affect the lipid contents in the pituitary anterior lobe (PAL) of adult male Wistar rats. As compared to metal non-exposed controls, PALs exposed to cadmium showed an increase in total phospholipid contents, which was associated to an increase of the incorporation of [1–¹⁴C]-methyl choline into phosphatidylcholine and of [U–¹⁴C]-glucose into total phospholipids. The incorporation of [1–¹⁴C]-methyl choline into sphingomyelin was not changed. Incorporation of [1–¹⁴C]-acetate into total fatty acids also increased but incorporation of [1–¹⁴C]-acetate into cholesterol did not change. The activity of phospholipase D decreased both in PALs from Cd exposed rats and in PAL dispersed cells treated with Cd in the culture medium from Cd non-exposed rats. In PALS from Cd exposed rats, a decrease of serum prolactin and growth hormone concentrations was determined. The results shown that cadmium modifies the lipid contents of pituitary gland and directly or indirectly the levels of prolactin and growth hormone in serum.     

Photoperiodic regulation of CS-ACS2, CS-ACS4 and CS-ERS gene expression contributes to the femaleness of cucumber flowers through diurnal ethylene production under short-day conditions

Photoperiod and the plant hormone, ethylene, modify sex expression of flowers in cucumber (Cucumis sativus L.). In the present study, femaleness of cucumber occurred under short‐day (8 h photoperiod) conditions compared to that under long‐day (16 h photoperiod) conditions, although the effect of photoperiod was more pronounced in a monoecious than in an andromonoecious cucumber. Application of ethylene had a greater effect than photoperiod on the production of female and bisexual flowers in monoecious and andromonoecious cucumbers, respectively. Ethylene evolution and the expression of CS‐ACS2, CS‐ACS4 and CS‐ERS genes in the shoot apices of both monoecious and andromonoecious cucumber plants had a diurnal rhythm with a peak in the middle of an 8 h or a 16 h light period. Peak ethylene evolution and expression of CS‐ACS2 was greater under short‐day conditions than under long‐day conditions in a monoecious cucumber but not in an andromonoecious one. Expression of CS‐ACS4 in monoecious and andromonoecious cucumber plants did not differ, but the level was higher under short‐day conditions compared with that under long‐day conditions. Thus, CS‐ACS2 and CS‐ACS4 might be involved in the basic diurnal rhythm of ethylene evolution in cucumber. Because exogenous ethylene increased the expression of CS‐ACS2 and CS‐ERS in monoecious cucumber possessing the M locus, but not in andromonoecious cucumber in which the function of the M locus was lost (Yamasaki et al. Plant and Cell Physiology 42, 608–619, 2001), the CS‐ACS2 gene might also be involved in ethylene production by positive feedback via regulation of M locus under short‐day conditions.     

Effects of physiological and abnormally elevated prolactin levels on the pituitary-testicular axis

Prolactin can influence testicular function both directly, and indirectly via altering release of gonadotropins from the pituitary. Although numerous effects of prolactin on male reproductive functions have been described, only a few were demonstrated in more than one species. These include effects on male accessory reproductive glands, on testicular luteinizing hormone receptors, on the release of gonadotropins and on sexual behavior. In rodents, prolactin appears to play a physiological role in the pituitary regulation of testicular function. This is especially pronounced in the golden hamster. In this seasonally-breeding species, alteration of prolactin release is one of the mechanisms mediating the effects of photoperiod on the testis. In the hamster, prolactin is required for the maintenance of luteinizing hormone and prolactin receptors in the testis and treatment with prolactin can completely reverse testicular atrophy induced by exposure to short photoperiod. In both men and rats, excessive release of prolactin (hyperprolactinemia) leads to suppression of sexual behavior and gonadotropin release. These effects appear to be due to the action of prolactin on the central nervous system. Most, if not all, of the effects of prolactin exhibit striking variability among species. Moreover, prolactin can exert differential effects on the same target tissue in the same species, depending primarily on the dose.     

Studies on human prolactin physiology

Although the clinical and experimental data were in favour of the existence of prolactin in humans like other vertebrates, as a pituitary hormone distinct from growth hormone, its presence remained contested until recent years. The predominant influence of the human hypothalamus on prolactin secretion is inhibitory. Circulating prolactin shows diurnal variations, which are not synchronized with that of TSH or ACTH; the prolactin rhythm is abolished during the last trimester of pregnancy and in patients with prolactin secreting tumors. Estrogens appeared to be less marked stimulators of prolactin secretion in man than in animals, although serum prolactin levels follow a pattern similar to that of endogenous estrogens during the normal menstrual cycle and during pregnancy. After delivery, basal prolactin levels declined progressively. In women under long term medroxyprogesterone acetate treatment, the immunoreactive serum prolactin was within the normal range of cycling women. Prolactin is found in appreciable amounts in amniotic fluid and in the serum of newborn infants. Synthetic LH and FSH releasing hormone did not change circulating prolactin levels in normal humans. A possible luteotrophic action of human prolactin in synergism with LH cannot be excluded.     

Seasonal prevalence and susceptibility to agrochemicals of Tyrophagus similis (Acari: Acaridae) in spinach buds and agricultural soil under greenhouse conditions

Seasonal prevalence of Tyrophagus similis was investigated from 1997 to 1998 in two spinach greenhouses in central Japan. Susceptibility of T. similis to agrochemicals was also tested in the laboratory. Tyrophagus similis density in the soil was low during the high temperature period from May to August. The density rapidly increased in late autumn and remained at a high level during the cool season from December to February. The number further increased in April and then rapidly decreased in May. The high temperatures in the greenhouses from spring to early autumn are considered the main causes of population decrease. Mites on spinach buds increased the number after those on and in the soil increased. Mites attacked spinach buds mostly in late autumn and early spring. Dichlorvos did not reduce the number of mites in either greenhouse even though it was highly toxic under laboratory condition. This discrepancy suggests that the mites in both the soil and spinach buds had little direct contact with the chemicals. These data suggest that once crop damage by mites is detected, it is usually too late to use chemicals, and that mites that live in the buds are protected from agrochemicals. This revised version was published online in July 2006 with corrections to the Cover Date.