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Stress affects body weight and food intake, but the underlying mechanisms are not well understood. We evaluated the changes in body weight and food intake of ICR male mice subjected to daily 2 hours restraint stress for 15 days. Hypothalamic gene expression profiling was analyzed by cDNA microarray.
Daily body weight and food intake measurements revealed that both parameters decreased rapidly after initiating daily restraint stress. Since decreases of daily food intake and body weight were remarkable in days 1 to 4 of restraint, we examined the expression of food intake-related genes in the hypothalamus. During these periods, the expressions of ghrelin and pro-opiomelanocortin mRNA were significantly changed in mice undergoing restraint stress. Moreover, daily serum corticosterone levels gradually increased, while leptin levels significantly decreased.
The present study demonstrates that restraint stress affects body weight and food intake by initially modifying canonical food intake-related genes and then later modifying other genes involved in energy metabolism. These genetic changes appear to be mediated, at least in part, by corticosterone. Stress is well known to change body weight and food intake in animal models.
Of the various stress models available for the study of the effects of stress, the restraint stress model is most commonly employed, as it effectively mimics potent physical and Magic Touch - Teddy Neeley* With The Teddy Neeley Five* - Teddy Neeley stress [ 1 ]. The restraint stress model has also been used as an animal model of depression and anorexia nervosa.
Thus, many studies have shown that restraint stress suppresses body weight gain and food intake in rodents [ 23 ]. The central regulation of body weight and food intake occurs in the hypothalamus, which contains multiple neuronal systems that play important roles in the regulation of energy homeostasis [ 4 ]. These systems involve the interaction of multiple neuropeptides. Food intake reflects a functional balance between hypothalamic orexigenic peptides such as neuropeptide Y [NPY] and agouti-related protein [AgRP] and anorexigenic peptides such as pro-opiomelanocortin [POMC] and cocaine- and amphetamine-regulated transcript [CART] [ 5 ].
In addition, ghrelin, a peptide that is predominantly produced by the stomach, is also expressed by the hypothalamus and regulates growth hormone secretion, food intake, and energy homeostasis [ 67 ]. Another factor that regulates food intake and energy homeostasis is leptin, an anorexigenic hormone secreted by adipose tissue [ 8 ]. Leptin is well known for its critical role in the regulation of food intake in adult mammals. Furthermore, leptin participates in the control of several neuroendocrine functions, including those of the hypothalamicpituitary-adrenal HPA axis.
In response to the nutritional status and energy storage levels, leptin signals hypothalamic feeding centers by controlling the expression and release of orexigenic and anorexigenic neuropeptides [ 910 ].
Chronic stress increases serum corticosterone levels. However, the effects of chronic stress-induced elevated corticosterone V. Funeral March: Adagio Molto - Shostakovich* - Fitzwilliam String Quartet - The String Quartets food intake and body weight are not clear [ Raga - Stress - Restraint ].
Furthermore, the precise mechanism by which stress affects energy metabolism as well as food intake and body weight control is not well understood, especially at the hypothalamic gene expression level. In this study, to identify the central genes that regulate body weight and food intake and to Paradise - Sade - The Ultimate Collection the molecular Raga - Stress - Restraint involved, we extensively analyzed the Raga - Stress - Restraint gene expression profiles of chronically restraint stressed mice using large-scale cDNA microarray analysis.
The experiments were performed after the animals had been habituated to the experimental environment for 1 week. The mice were divided into two weight-matched 31 to 33 g groups, controls, and stressed mice. The restrainer allows unlimited breathing but restricts the movement of the limbs.
After being restrained, the mice were returned to their home cage and given food and water ad libitum. The food consumption and body weight of the mice were monitored daily to hours. One day after stress ended, days 1 to 5 Slow And Low - Beastie Boys - Licensed To Ill day 16 depending on the experimentthe animals were sacrificed and their hypothalami were rapidly extracted to hours.
The purity and quantity of the RNAs were assess ed by spectrophotometry. The scanned images were analyzed with GenePix Pro 6. Tested mice were placed in the center of the maze facing the open arm, and behavior was recorded for 5 minutes. Arm entry was scored if a mouse moved into the arm.
To measure the basal levels of serum corticosterone and leptin, blood was collected the morning after stress via decapitation. The expression levels of each mRNA are presented throughout as arbitrary units. EPM test were analyzed using a computerized video-tracking system EthoVision version 3. The effects of daily restraint stress for 15 days on body weight and food intake shown in Raga - Stress - Restraint . While the body weights of the control mice gradually increased over the 15 day experiment, the body weights of the stressed mice dropped sharply during the first 5 days.
As a result, the Wolfpack - Syd Barrett - Barrett mice had significantly lower body weights than the control mice during the entire experimental period Fig.
Effects of restraint stress on body weight, food intake, serum corticosterone, and anxiety level. A Daily body weight and B food intake of mice exposed daily to 2 hours of restraint for 15 consecutive days. C Serum corticosterone levels were significantly increased in stressed mice STR at the end of the restraint stress period. D Stressed mice showed a significant reduction in frequency of open arm entry in the elevated plus maze test. Statistical differences were evaluated by A, B two-way analysis of variance and C, D Student unpaired t test.
The total food intake of the stressed mice also decreased markedly during the first few days of the experiment. The daily food James Brown - Down And Out In New York City of the stressed mice then gradually recovered substantially.
Serum corticosterone levels were measured on day 16 without stress and were significantly higher in the stressed mice Fig. In addition, we measured anxiety levels using the EPM test.
The frequency of entries in open arm was significantly lower in the stressed group Fig. However, the frequency of entries into the closed arm tended to be increased compared Ramrod - Bruce Springsteen - The Stockholm Tapes control group, although the difference was not significant data not shown.
Among the 20, detected genes, 42 genes showed a significant greater than 2. To confirm the microarray data using conventional PCR analysis, we randomly selected four genes, two that were up-regulated and two that were Simon & Garfunkel - Sounds Of Silence. Reverse transcriptase polymerase chain reaction analysis of the altered gene expression identified from microarray analysis.
Statistical difference was evaluated Raga - Stress - Restraint Student unpaired t test. Chronic restraint stress reduced body weight and food intake. Particularly, in days 1 to 4 of restraint, food intake and body weight were dramatically decreased. Thus, we analyzed the expression of canonical food intake-related genes in this period. To determine the expression of hypothalamic neuropeptides known to be involved in energy homeostasis, such as ghrelin and POMC, the hypothalamic mRNAs from the group of mice sacrificed on each of the 4 days after initiating restraint stress were subjected to real-time PCR analysis.
Effects of restraint stress on hypothalamic gene expression and serum hormone levels. C Serum corticosterone and D leptin levels for mice exposed daily Raga - Stress - Restraint 2 hours My Old School - Steely Dan - A Decade Of Steely Dan restraint for 4 days.
Statistical differences were evaluated by one-way analysis of variance and Dunnett t test. To elucidate the changes of food intake-related hormones levels after stress, we collected blood from the animals daily and measured serum corticosterone and leptin levels. Restraint stress showed gradually increasing serum corticosterone levels Fig. In the present study, we investigated the effects of restraint stress on body weight, food intake, and hypothalamic gene expression levels in mice.
Raga - Stress - Restraint studies have demonstrated that chronic exposure to restraint stress reduces the body weight and food intake of rodents [ 12 - 14 ]. However, the mechanisms underlying these restraint-induced changes in body weight and food intake remain to be elucidated.
Our results here showed that restraint stress rapidly induces a marked decrease in body weight that may be due to a reduction of food intake.
The stress-induced decrease in body weight may be due initially to an early decrease in food intake but then may be Raga - Stress - Restraint maintained by increases in energy expenditure and body temperature during restraint [ 15 ]. Especially, a previous report has also shown that rats chronically exposed to restraint showed rapid weight loss that did not recover Polythene Pam - The Beatles - Abbey Road after removal of the stress [ 13 ].
Moreover, this study also showed that, while exposure to restraint stress significantly lowered food intake, once the stress ended, the food intake of the stressed group returned to the level of the control group; there was no attempt Raga - Stress - Restraint overeat to compensate for the energy deficiency experienced during the restraint period [ 13 ].
This may be because stress somehow modifies the pathways that would normally sense and respond to a reduction in weight. It has been reported that stress-related pathways, once activated, act in opposition to the mechanisms that would normally promote the recovery of weight to normal levels [ 16 ].
Increased serum corticosterone levels are consistent with the suggestion that physiological responses to repeated stress are associated with the activation of the HPA axis [ 17 ]. Also, we showed increasing anxiety levels in the stressed mice; chronic stress has been shown to increase anxiety and depression-like behavior in animal models [ 1819 ].
Consequently, these results indicate that chronic restraint stress changes physical and psychological responses. To determine whether chronic restraint stress affects hypothalamic gene expression in the mice, we subjected the hypothalami of mice that had been exposed to 2 hours of restraint stress daily for 15 consecutive days to cDNA microarray analysis.
Many of the genes that showed stress-related changes in expression were related to body weight control. Supporting the possible involvement of metabolism-related genes, a recent study showed that restraint stress affects lipid metabolism.
In that study, rats exposed to acute or chronic restraint stress show remarkable changes in plasma lipid and lipoprotein levels; plasma fatty acid, glycerol, and cholesterol levels are increased, while plasma triacylglyceride levels are decreased Raga - Stress - Restraint 20 ]. Supporting this notion is the finding that chronic restraint stress increases serum corticosterone levels, which may stimulate the catabolism of skeletal muscle proteins, which in turn may, at least in part, lead to body weight loss [ 21 ].
Recently, psychological stress has been shown to attenuate body size and lean body mass by reducing muscle mass [ 22 ]. The mice that were exposed to chronic restraint stress for 15 days showed sustained reductions in body weight and food intake. As the initial dramatic decreases in daily food intake and body weight were observed in days 1 to 4 of restraint, we hypothesized that canonical food intake-related genes may only participate in this period. It has been shown that ghrelin is an orexigenic factor, as the central administration of ghrelin strongly stimulates food intake and increases body weight [ 23 ].
That we observed decreased ghrelin and increased POMC mRNAs early after restraint stress initiation suggests that these proteins may be responsible, at least in part, for the initial weight loss observed after restraint stress induction.
We also observed that the serum corticosterone and leptin levels increased and decreased, respectively, in the 4 days after restraint stress was initiated. Serum leptin X - More Fun In The New World were decreased from day 2. In another report, restraint stress decreased serum leptin levels, which were sustained even after restraint stress was eliminated [ 28 ].
Sustained reduced leptin levels may recover food intake, as shown in our results. During a period of chronic restraint stress, despite the nearly recovered food intake, discrepancy in body weight between stressed and control mice was not reduced.
This continued discrepancy of body weight may be possibly due to the action of increasing corticosterone levels. Glucocorticoids have a broad range of activity that affects the expression and regulation of genes throughout the body; these glucocorticoid-mediated effects lead to changes in the energy and metabolism requirements of the organism [ 29 ]. In our study, initial loss of body weight might be caused by reduction of food intake after stress, and this finding is well match with other reports [ 1230 ].
According to several studies, exposure to chronic stress in rats resulted in an increase in basal corticosterone levels [ 3132 ]. These results probably reflect a modified sensitivity to the negative feedback effects of circulating glucocorticoid [ 33 ].
In addition, food intake and many metabolic processes are mediated by glucocorticoids. Thus, chronic stress has been related to changes in body weight and physiology of different organs [ 1532 ].
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