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type=\u0022text\/css\u0022 rel=\u0022stylesheet\u0022 href=\u0022\/\/d282kpwvnogo5m.cloudfront.net\/sites\/default\/files\/cdn\/css\/http\/css_Xg7z6oCTVgud_Q0huYz9x9iiD5H_2YPSJ5z2ZViSWdY.css\u0022 media=\u0022all\u0022 \/\u003E\n\u003Clink rel=\u0027stylesheet\u0027 type=\u0027text\/css\u0027 href=\u0027\/sites\/all\/modules\/contrib\/panels\/plugins\/layouts\/onecol\/onecol.css\u0027 \/\u003E\u003C\/head\u003E\u003Cbody\u003E\u003Cdiv class=\u0022panels-ajax-tab-panel panels-ajax-tab-panel-sageoa-tab-art\u0022\u003E\u003Cdiv class=\u0022panel-display panel-1col clearfix\u0022 \u003E\n  \u003Cdiv class=\u0022panel-panel panel-col\u0022\u003E\n    \u003Cdiv\u003E\u003Cdiv class=\u0022panel-pane pane-highwire-markup\u0022 \u003E\n  \n      \n  \n  \u003Cdiv class=\u0022pane-content\u0022\u003E\n    \u003Cdiv class=\u0022highwire-markup\u0022\u003E\u003Cdiv xmlns=\u0022http:\/\/www.w3.org\/1999\/xhtml\u0022 id=\u0022content-block-markup\u0022 xmlns:xhtml=\u0022http:\/\/www.w3.org\/1999\/xhtml\u0022\u003E\u003Cdiv class=\u0022article fulltext-view \u0022\u003E\u003Cspan class=\u0022highwire-journal-article-marker-start\u0022\u003E\u003C\/span\u003E\u003Cdiv class=\u0022section abstract\u0022 id=\u0022abstract-1\u0022\u003E\u003Ch2\u003ESummary\u003C\/h2\u003E\n            \u003Cp id=\u0022p-1\u0022\u003ENumerous recent studies of animal models suggest a highly active role for the central nervous system in energy homeostasis. While this is not surprising, an expansion of these findings indicates that insulin signaling has a pronounced influence on the lifespan of a wide array of organisms.\u003C\/p\u003E\n         \u003C\/div\u003E\u003Cul class=\u0022kwd-group\u0022\u003E\u003Cli class=\u0022kwd\u0022\u003Einsulin\u003C\/li\u003E\u003Cli class=\u0022kwd\u0022\u003Eendocrinology\u003C\/li\u003E\u003C\/ul\u003E\u003Cdiv class=\u0022section\u0022 id=\u0022sec-1\u0022\u003E\n         \n         \u003Cp id=\u0022p-2\u0022\u003ENumerous recent studies of animal models suggest a highly active role for the central nervous system (CNS) in energy homeostasis. While this is not surprising, an expansion of these findings indicates that insulin signaling has a pronounced influence on the lifespan of a wide array of organisms. Jens Claus Br\u00fcning, MD, University of Cologne, Cologne, Germany, and winner of the 2008 Oskar Minkowski Prize, discussed these data in the 43\u003Csup\u003Erd\u003C\/sup\u003E Minkowski Lecture.\u003C\/p\u003E\n         \u003Cp id=\u0022p-3\u0022\u003ETo the frustration of some researchers, insulin has been shown to have pleiotropic effects in nearly all tissues. Yet to date, the primary focus of these investigations has been on insulin activity in the peripheral tissues. Indeed, for many years insulin also was the primary hormone, or signaling molecule under consideration, but there was a paradigm shift in the understanding of metabolic regulation with the discovery and eventual isolation of leptin.\u003C\/p\u003E\n         \u003Cp id=\u0022p-4\u0022\u003ELeptin created a sensation in the lay press as well as in the peer-reviewed literature, because its activity in the body affected feeding behavior. Mice with defective leptin expression were observed to be obese (Shang et al. \u003Cem\u003ENature\u003C\/em\u003E 1994). Leptin deficiency in humans also was identified and fueled the assumption that the cause of obesity had indeed been found. Although patients did respond dramatically to leptin augmentation, the underlying hormonal defect was determined to be fairly rare. Research then shifted in perspective to consider not only the signal, but also the receiver. Leptin that is produced in adipose tissue has been shown to bind to the ventromedial nucleus of the hypothalamus, the center of appetite control. The message that is carried by this diminutive 16-kD protein is that the body has met its energy requirements, stimulating the brain to send signals to the body to stop eating.\u003C\/p\u003E\n         \u003Cp id=\u0022p-5\u0022\u003EThis activity was further localized to the ARC (hypothalamic arcuate nucleus) neurons, and thereafter multiple signals were identified as being targeted to this area. These included leptin, insulin, GLP-1, and glucose (\u003Ca id=\u0022xref-fig-1-1\u0022 class=\u0022xref-fig\u0022 href=\u0022#F1\u0022\u003EFigure 1\u003C\/a\u003E). The focus on ARC neurons was further validated by murine investigations, showing that if agouti-related peptide and neuropeptide Y, normally expressed in ARC, were disabled, feeding behavior was unaltered. However, if the neurons that expressed these genes were themselves ablated, food intake was drastically reduced, nearly to the point of starvation. ARC neurons are now known to play a critical role in the regulation of energy homeostasis (Gropp et al. \u003Cem\u003ENat Neuro\u003C\/em\u003E 2005).\u003C\/p\u003E\n         \u003Cdiv id=\u0022F1\u0022 class=\u0022fig pos-float  odd\u0022\u003E\u003Cdiv class=\u0022highwire-figure\u0022\u003E\u003Cdiv class=\u0022fig-inline-img-wrapper\u0022\u003E\u003Cdiv class=\u0022fig-inline-img\u0022\u003E\u003Ca href=\u0022http:\/\/d282kpwvnogo5m.cloudfront.net\/content\/spmdc\/8\/7\/31\/F1.large.jpg?width=800\u0026amp;height=600\u0026amp;carousel=1\u0022 title=\u0022ARC Neurons as Targets of Multiple Signals.\u0022 class=\u0022fragment-images colorbox-load\u0022 rel=\u0022gallery-fragment-images-1029255666\u0022 data-figure-caption=\u0022ARC Neurons as Targets of Multiple Signals.\u0022 data-icon-position=\u0022\u0022 data-hide-link-title=\u00220\u0022\u003E\u003Cimg class=\u0022fragment-image\u0022 alt=\u0022Figure 1.\u0022 src=\u0022http:\/\/d282kpwvnogo5m.cloudfront.net\/content\/spmdc\/8\/7\/31\/F1.medium.gif\u0022\/\u003E\u003C\/a\u003E\u003C\/div\u003E\u003C\/div\u003E\u003Cul class=\u0022highwire-figure-links inline\u0022\u003E\u003Cli class=\u00220 first\u0022\u003E\u003Ca href=\u0022http:\/\/d282kpwvnogo5m.cloudfront.net\/content\/spmdc\/8\/7\/31\/F1.large.jpg?download=true\u0022 class=\u0022highwire-figure-link highwire-figure-link-download\u0022 title=\u0022Download Figure 1.\u0022 data-icon-position=\u0022\u0022 data-hide-link-title=\u00220\u0022\u003EDownload figure\u003C\/a\u003E\u003C\/li\u003E\u003Cli class=\u00221\u0022\u003E\u003Ca href=\u0022http:\/\/d282kpwvnogo5m.cloudfront.net\/content\/spmdc\/8\/7\/31\/F1.large.jpg\u0022 class=\u0022highwire-figure-link highwire-figure-link-newtab\u0022 target=\u0022_blank\u0022 data-icon-position=\u0022\u0022 data-hide-link-title=\u00220\u0022\u003EOpen in new tab\u003C\/a\u003E\u003C\/li\u003E\u003Cli class=\u00222 last\u0022\u003E\u003Ca href=\u0022\/highwire\/powerpoint\/11293\u0022 class=\u0022highwire-figure-link highwire-figure-link-ppt\u0022 data-icon-position=\u0022\u0022 data-hide-link-title=\u00220\u0022\u003EDownload powerpoint\u003C\/a\u003E\u003C\/li\u003E\u003C\/ul\u003E\u003C\/div\u003E\u003Cdiv class=\u0022fig-caption\u0022 xmlns:xhtml=\u0022http:\/\/www.w3.org\/1999\/xhtml\u0022\u003E\u003Cspan class=\u0022fig-label\u0022\u003EFigure 1.\u003C\/span\u003E \n               \u003Cp id=\u0022p-6\u0022 class=\u0022first-child\u0022\u003EARC Neurons as Targets of Multiple Signals.\u003C\/p\u003E\n            \u003Cdiv class=\u0022sb-div caption-clear\u0022\u003E\u003C\/div\u003E\u003C\/div\u003E\u003C\/div\u003E\n         \u003Cp id=\u0022p-7\u0022\u003EInsulin signaling in peripheral tissues entails activation of the insulin receptor substrate (IRS)-phosphatidylinositol 3-kinase (PI3K) enzyme system. In the hypothalamus, insulin functions with leptin as an afferent adiposity signal that is important for the regulation of body fat stores and hepatic glucose metabolism. Studies have shown that the IRS-PI3K pathway is a mediator of insulin action in the arcuate nucleus as well. Combined with recent evidence that leptin activates PI3K signaling in the hypothalamus, this suggests a conduit for neuronal crosstalk between insulin and leptin signaling (Niswender et al. \u003Cem\u003EDiabetes\u003C\/em\u003E 2003).\u003C\/p\u003E\n         \u003Cp id=\u0022p-8\u0022\u003EMice that lack IR have proven to be invaluable in gaining a better understanding of the role of insulin action in the brain. Downregulation of neuronal IR expression results in severe hyperinsulinemia and a dramatic upregulation of hepatic leptin receptor expression. Leptin replacement restored normal glucose metabolism in these mice. Insulin action, specifically in AgRP-expressing neurons, does play a critical role in controlling hepatic glucose production and may provide a target for the treatment of insulin resistance in type 2 diabetes (Koch et al. \u003Cem\u003EJCI\u003C\/em\u003E 2008; Konner et al. \u003Cem\u003ECell Metab\u003C\/em\u003E 2007).\u003C\/p\u003E\n         \u003Cp id=\u0022p-9\u0022\u003ECaloric restriction has been strongly associated with a reduction in aging-associated pathologies, thereby resulting in a longer, healthier life. As first demonstrated in \u003Cem\u003EDrosophila\u003C\/em\u003E (Chapman et al. \u003Cem\u003EProc Roy Soc\u003C\/em\u003E 1996), and later in \u003Cem\u003EC. elegans\u003C\/em\u003E and mice, it is clear that restricted caloric intake can increase longevity.\u003C\/p\u003E\n         \u003Cp id=\u0022p-10\u0022\u003ESelman et al. were able to localize this benefit to the CNS, observing that IRS-1-deficient mice stay healthy and disease-free longer and had the ability to maintain glucose homeostasis (FASEB 2008). This work was replicated in \u003Cem\u003EC. elegans,\u003C\/em\u003E verifying that the nervous system\u0027s relationship with insulin is a central regulator of animal longevity (Wolkow et al. \u003Cem\u003EScience\u003C\/em\u003E 2000; \u003Ca id=\u0022xref-fig-2-1\u0022 class=\u0022xref-fig\u0022 href=\u0022#F2\u0022\u003EFigure 2\u003C\/a\u003E).\u003C\/p\u003E\n         \u003Cp id=\u0022p-11\u0022\u003EThe residence of such activity in the CNS raises a question about whether glucose dysregulation plays a role in age-related diseases of the brain, such as Alzheimer disease (AD). An animal model of AD was shown to have downregulated IGF-1R and IR and an excessive presence of their key substrate adaptor proteins, IRS-1 and IRS-2, the physiological equivalent of insulin resistance. The increase in IGF-1R was detected around and within amyloid beta-containing plaques, the pathological hallmark of AD, and resistant to IGF-1R\/IR signaling (Moloney et al. \u003Cem\u003ENeurobiol Aging\u003C\/em\u003E 2008); yet, this link is merely hypothesis-generating. One recent study by Moroz et al. assessed AD-type neurodegeneration in a type 2 diabetes mellitus (T2DM) mouse model but did not observe an AD histopathology, leading the authors to conclude that obesity and T2DM may contribute, but are not sufficient, to cause AD (Moroz et al. \u003Cem\u003EJ Alzheimer\u0027s Disease\u003C\/em\u003E 2008).\u003C\/p\u003E\n         \u003Cdiv id=\u0022F2\u0022 class=\u0022fig pos-float  odd\u0022\u003E\u003Cdiv class=\u0022highwire-figure\u0022\u003E\u003Cdiv class=\u0022fig-inline-img-wrapper\u0022\u003E\u003Cdiv class=\u0022fig-inline-img\u0022\u003E\u003Ca href=\u0022http:\/\/d282kpwvnogo5m.cloudfront.net\/content\/spmdc\/8\/7\/31\/F2.large.jpg?width=800\u0026amp;height=600\u0026amp;carousel=1\u0022 title=\u0022Restoring IIS in the CNS Abrogates the Life-Extending Effect of Systemic IR in C elegans.\u0022 class=\u0022fragment-images colorbox-load\u0022 rel=\u0022gallery-fragment-images-1029255666\u0022 data-figure-caption=\u0022\u0026amp;lt;div xmlns=\u0026amp;quot;http:\/\/www.w3.org\/1999\/xhtml\u0026amp;quot;\u0026amp;gt;Restoring IIS in the CNS Abrogates the Life-Extending Effect of Systemic IR in \u0026amp;lt;em\u0026amp;gt;C elegans\u0026amp;lt;\/em\u0026amp;gt;.\u0026amp;lt;\/div\u0026amp;gt;\u0022 data-icon-position=\u0022\u0022 data-hide-link-title=\u00220\u0022\u003E\u003Cimg class=\u0022fragment-image\u0022 alt=\u0022Figure 2.\u0022 src=\u0022http:\/\/d282kpwvnogo5m.cloudfront.net\/content\/spmdc\/8\/7\/31\/F2.medium.gif\u0022\/\u003E\u003C\/a\u003E\u003C\/div\u003E\u003C\/div\u003E\u003Cul class=\u0022highwire-figure-links inline\u0022\u003E\u003Cli class=\u00220 first\u0022\u003E\u003Ca href=\u0022http:\/\/d282kpwvnogo5m.cloudfront.net\/content\/spmdc\/8\/7\/31\/F2.large.jpg?download=true\u0022 class=\u0022highwire-figure-link highwire-figure-link-download\u0022 title=\u0022Download Figure 2.\u0022 data-icon-position=\u0022\u0022 data-hide-link-title=\u00220\u0022\u003EDownload figure\u003C\/a\u003E\u003C\/li\u003E\u003Cli class=\u00221\u0022\u003E\u003Ca href=\u0022http:\/\/d282kpwvnogo5m.cloudfront.net\/content\/spmdc\/8\/7\/31\/F2.large.jpg\u0022 class=\u0022highwire-figure-link highwire-figure-link-newtab\u0022 target=\u0022_blank\u0022 data-icon-position=\u0022\u0022 data-hide-link-title=\u00220\u0022\u003EOpen in new tab\u003C\/a\u003E\u003C\/li\u003E\u003Cli class=\u00222 last\u0022\u003E\u003Ca href=\u0022\/highwire\/powerpoint\/11294\u0022 class=\u0022highwire-figure-link highwire-figure-link-ppt\u0022 data-icon-position=\u0022\u0022 data-hide-link-title=\u00220\u0022\u003EDownload powerpoint\u003C\/a\u003E\u003C\/li\u003E\u003C\/ul\u003E\u003C\/div\u003E\u003Cdiv class=\u0022fig-caption\u0022\u003E\u003Cspan class=\u0022fig-label\u0022\u003EFigure 2.\u003C\/span\u003E \n               \u003Cp id=\u0022p-12\u0022 class=\u0022first-child\u0022\u003ERestoring IIS in the CNS Abrogates the Life-Extending Effect of Systemic IR in \u003Cem\u003EC elegans\u003C\/em\u003E.\u003C\/p\u003E\n            \u003Cdiv class=\u0022sb-div caption-clear\u0022\u003E\u003C\/div\u003E\u003C\/div\u003E\u003C\/div\u003E\n         \u003Cp id=\u0022p-13\u0022\u003EThe physiological importance of this homeostatic control system is highlighted by the severe obesity that results from the dysfunction of any of its key components. This new information provides a biological context within which to consider the global obesity epidemic and identifies numerous potential avenues for therapeutic intervention and future research (Schwartz et al. \u003Cem\u003ENature\u003C\/em\u003E 2006).\u003C\/p\u003E\n      \u003C\/div\u003E\u003Cul class=\u0022copyright-statement\u0022\u003E\u003Cli class=\u0022fn\u0022 id=\u0022copyright-statement-1\u0022\u003E\u00a9 2008 MD Conference Express\u003C\/li\u003E\u003C\/ul\u003E\u003Cspan class=\u0022highwire-journal-article-marker-end\u0022\u003E\u003C\/span\u003E\u003C\/div\u003E\u003Cspan id=\u0022related-urls\u0022\u003E\u003C\/span\u003E\u003C\/div\u003E\u003Ca href=\u0022http:\/\/mdc.sagepub.com\/content\/8\/7\/31.abstract\u0022 class=\u0022hw-link hw-link-article-abstract\u0022 data-icon-position=\u0022\u0022 data-hide-link-title=\u00220\u0022\u003EView Summary\u003C\/a\u003E\u003C\/div\u003E  \u003C\/div\u003E\n\n  \n  \u003C\/div\u003E\n\u003C\/div\u003E\n  \u003C\/div\u003E\n\u003C\/div\u003E\n\u003C\/div\u003E\u003Cscript type=\u0022text\/javascript\u0022 src=\u0022http:\/\/mdc.sagepub.com\/sites\/all\/modules\/highwire\/highwire\/plugins\/highwire_markup_process\/js\/highwire_figures.js?nzmcjp\u0022\u003E\u003C\/script\u003E\n\u003Cscript type=\u0022text\/javascript\u0022 src=\u0022http:\/\/mdc.sagepub.com\/sites\/all\/modules\/highwire\/highwire\/plugins\/highwire_markup_process\/js\/highwire_openurl.js?nzmcjp\u0022\u003E\u003C\/script\u003E\n\u003C\/body\u003E\u003C\/html\u003E"}