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A schematic drawing outlining the main signaling pathways activated in resp=
onse to ligand engagement with the GLP-1R and their major downstream effect=
s on acute insulin secretion, insulin synthesis, preservation of =CE=B2 cel=
l function and mass and regulation of proliferation. Pathways are glucose d=
ependent hence the inclusion of glucose metabolism. GLP-1/Ex-4 bind to GLP-=
1R causing an increase in cAMP (Drucker et al., 1987); this leads to activa=
tion of both PKA (Wang et al., 2001) and EPAC (Holz, 2004). Localized low c=
oncentrations of cAMP lead to preferential activation of PKA. Higher cell-w=
ide increases of cAMP by the AC stimulator forskolin (FSK) or the phosphodi=
esterase (PDE) inhibitor IBMX favor the EPAC pathway. cAMP is compartmental=
ized by PDEs most notably the PDE3B isoform as shown (Harndahl et al., 2004=
). PKA anchoring proteins (AKAPs) influence the specificity of cAMP respons=
e by anchoring the PKA to specific intracellular sites (Lester et al., 1997=
). Shown here also is the Ca2+/calmodulin binding protein IQGAP1 which co-i=
mmunoprecipitates with PKA and AKAP79 (Nauert et al., 2003). cAMP levels ar=
e increased as a consequence of ATP activation of AC consequent upon glucos=
e metabolism. Binding of cAMP to the regulatory units of PKA results in rel=
ease of the catalytic units from PKA and its activation. Sustained oscillat=
ory increases in cAMP by GLP-1R activation lead to translocation of PKA to =
the nucleus (Dyachok et al., 2006; Gao et al., 2002) where it regulates PDX=
-1 (Wang et al., 2001) and CREB activation and subsequently insulin transcr=
iption (Chepurny et al., 2002; Hay et al., 2005; Kemp and Habener, 2001). D=
ownstream targets of PKA and Epac in acute insulin secretion, include the K=
ATP and Kv channels, the insulin secretory vesicles and the IP3 Ca2+ channe=
ls on the endoplasmic reticulum (ER). P38 MAPK, although activated by GLP-1=
R agonists (Kemp and Habener, 2001; Montrose-Rafizadeh et al., 1999) is not=
 included as the exact mechanism of activation has not been described. Acti=
vation of the MEK/ERK pathway is coordinated through both the Epac moieties=
 and the Ca2+/calmodulin kinases (Arnette et al., 2003; Gomez et al., 2002)=
. The effect of PKA on CREB mediated induction of the IRS2 gene is shown, t=
his is a prolonged effect of GLP-1R activation (Jhala et al., 2003). Acutel=
y PI3 kinase is also stimulated by transactivation of the EGF receptor by c=
Src-activated betacellulin (BTC; Buteau et al., 2003). Downstream of PI3 ki=
nase are PKB and PKC=CE=B6 both of which are implicated in =CE=B2 cell prol=
iferation and PKB in prevention of =CE=B2 cell death (Buteau et al., 2001; =
Wang and Brubaker, 2002). FoxO1 is regulated by phosphorylation by PKB whic=
h results in its exclusion from the nucleus thus permitting the nuclear tra=
nslocation of PDX-1 (Buteau et al., 2006). Finally enhanced ATP production =
due to increased mobilization of Ca2+ which in turn upregulate mitochondria=
l dehydrogenases leads to upregulation of mTOR activity and its downstream =
effector S6K1 (Kwon et al., 2004a). mTOR is implicated in increased =CE=B2 =
cell mitosis and may also be activated by PKB. GLP-1R activation also leads=
 to stabilization of the insulin transcript by stimulating nucleocytoplasmi=
c translocation of polypyrimidine tract binding protein (PTB) which binds t=
o the U-rich polypyrimidine tract of insulin and insulin secretory vesicle =
mRNA transcripts thereby stabilizing them (Knoch et al., 2006). Mechanisms =
that have not been clearly demonstrated are shown by broken arrows.



Simplified schema of the human (A) and rat I (B) insulin promoters. The ele=
ments known to be regulated downstream of GLP-1R activation are shown in bl=
ue. There are four CRE sites in the insulin gene two upstream (CRE1 and CRE=
2) and two downstream (CRE3 and CRE4) of the transcription start site. With=
 the exception of the first one (CRE1) all were shown to be induced when co=
nstructs containing fragments representing the individual sites were transf=
ected into INS-1 cells (Hay et al., 2005). It is probable that the close pr=
oximity of the CRE1 site to the A3 element to which PDX-1 binds actively pr=
ohibits complex formation at this CRE site. All of the NFAT sites studied i=
n the rat I insulin promoter are responsive to the combination of glucose a=
nd GLP-1, although NFAT2 was relatively insensitive to GLP-1 alone (Lawrenc=
e et al., 2002)

Schema outlining the signaling mechanisms reported to be involved in GLP-1R=
-induced differentiation/neogenesis of pancreatic precursor cells, prolifer=
ation and in the prevention of apoptosis. Dashed lines indicate mechanisms =
that are either not fully delineated or in the case of the cAMP activation =
of the MEK pathway are complex and are shown completely in Fig 2. The mecha=
nism shown for involvement of BMP and TGF=CE=B2 signaling pathways in diffe=
rentiation is after Gittes and co-workers (Yew et al., 2005; Yew et al., 20=
04). PKB is shown as inhibiting FoxO1 (by phosphorylation). When FoxO1 is i=
n its active and unphosphorylated state it inhibits the transcription of PD=
X-1 and its translocation to the nucleus (Kitamura et al., 2002).

Wang F, Zhang XL, Zhou Y, Ye L, Qi Z and Wu J (2005) Type IVB piliated Salm=
onella typhi enhance IL-6 and NF-kB production in human monocytic THP-1 cel=
ls through activation of protein kinase C. Immunobiology 210 283-293

Wang GP, Qi ZH and Chen FP (2005) Advances of study on activation of nuclea=
r factor kappaB in hematological malignancies and its mechanism [Article in=
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Wang HJ, Wu ZY, Fan P and Bian JM (2005) The nuclear factor kappa B activat=
ion: the key step of cell proliferation in estrogen receptor-negative breas=
t cancer cells [Article in Chinese]. Zhonghua Wai Ke Za Zhi 43 1014-1016

Wang J, Ouyang W, Li J, Wei L, Ma Q, Zhang Z, Tong Q, He J and Huang C (200=
5) Loss of tumor suppressor p53 decreases PTEN expression and enhances sign=
aling pathways leading to activation of activator protein 1 and nuclear fac=
tor kB induced by UV radiation. Cancer Research 65 6601-6611

Wang K, Brems JJ, Gamelli RL and Ding J (2005) Reversibility of caspase act=
ivation and its role during glycochenodeoxycholate-induced hepatocyte apopt=
osis. Journal of Biological Chemistry 280 23490-23495

Wang SY, Feng R, Bowman L, Penhallegon R, Ding M and Lu Y (2005) Antioxidan=
t activity in Lingonberries (Vaccinium vitis-idaea L.) and its inhibitory e=
ffect on activator protein-1, nuclear factor-kB, and mitogen-activated prot=
ein kinases activation. J Agric Food Chem 53 3156-3166

Wang T, Hu YC, Dong S, Fan M, Tamae D, Ozeki M, Gao Q, Gius D and Li JJ (20=
05) Co-activation of ERK, NF-kB and GADD45beta in response to ionizing radi=
ation. Journal of Biological Chemistr y 280 12593-12601

Wang YH, Wang WY, Chang CC, Liou KT, Sung YJ, Liao JF, Chen CF, Chang S, Ho=
u YC, Chou YC and Shen YC (2005) Taxifolin ameliorates cerebral ischemia-re=
perfusion injury in rats through its anti-oxidative effect and modulation o=
f NF-kB activation. Journal of Biomedical Science 13 1127-141

Ward C, Murray J, Clugston A, Dransfield I, Haslett C and Rossi AG (2005) I=
nterleukin-10 inhibits lipopolysaccharide-induced survival and extracellula=
r signal-regulated kinase activation in human neutrophils. European Journal=
 of Immunolog y 35 2728-2737

Watson JL, Vicario M, Wang A, Moreto M and McKay DM (2005) Immune cell acti=
vation and subsequent epithelial dysfunction by Staphylococcus enterotoxin =
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ellular Immunology 237 7-16

Wei CY, Huang KC, Chou YH, Hsieh PF, Lin KH and Lin WW (2005) The role of R=
ho-associated kinase in differential regulation by statins of interleukin-1=
beta and lipopolysaccharide-mediated NF-kB activation and inducible nitric =
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harmacology 69 960-967

Wei JF, Sun K, Xu SG, Xie HY and Zheng SS (2005) Inhibition of PMA-induced =
endothelial cell activation and adhesion by over-expression of domain negat=
ive IkBa protein. World Journal of Gastroenterology 11 3080-3084

Werber S, Shalit I, Fabian I, Steuer G, Weiss T and Blau H (2005) Moxifloxa=
cin inhibits cytokine-induced MAP kinase and NF-kB activation as well as ni=
tric oxide synthesis in a human respiratory epithelial cell line. Journal o=
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