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Simply put, it seems that the life expectancy of the mice allows expression of only the most rapidly growing tumor types. The situation for the development of anterior pituitary tumors is similar, in which PRLomas are the most commonly occurring tumor in MEN1 patients, who occasionally develop somatotrophinomas and rarely corticotrophinomas.

The observed higher proliferation rates of lactotrophs and the mathematical modeling analysis Figs. Thus, the mathematical modeling of anterior pituitary NET growth has provided insights into the variety of pituitary NET types that are found and suggests that pituitary lactotrophs and somatotrophs are most susceptible to tumorigenesis from loss of the wild-type Men1 allele Fig.

However, the higher proportion of lactotrophs and somatotrophs in the anterior pituitary, when compared with corticotrophs, gonadotrophs, and thyrotrophs, may also contribute to the higher prevalence of lactotrophinomas and somatotrophinomas.

These in silico predictions correlate with observations in MEN1 patients and Men1 mouse models in which insulinomas and lactotrophinomas frequently arise as the pancreatic and pituitary NETs, respectively 13 , However, it is important to note the limitations of the mathematical modeling, which does not consider the occurrence of additional mutations in genes other than MEN1 that may suddenly increase the proliferation rates and conferment of invasive and malignant potential, and the changes in tumor hormonal expression during growth that may render a nonsecreting tumor into a secreting tumor, or vice versa.

The mathematical model Fig. Our mathematical modeling, which demonstrates the importance of the occurrence of mutations in both Men1 alleles Fig.

This prediction could be tested by breeding conditional Men1 knockout mice with appropriate transgenic mice expressing Cre under the control of a tissue-specific promoter Thus, development of glucagonomas may be related to the occurrence of the Men1 mutations, and use of other tissue-specific conditional Men1 knockout mice may help to further assess the validity of the mathematical modeling and the development of other NETs in the pancreas and pituitary.

The apoptotic response was found to differ in the endocrine tumors. Thus, apoptosis was increased in the gonadal tumors Fig.

These findings are difficult to reconcile with a unifying explanation. Menin has been reported, by in vitro studies, to promote the transcription of caspase 8 and stimulate the extrinsic apoptotic pathway 12 , 49 , However, the findings of increased apoptosis in the gonadal tumors and no change in the pancreatic NETs, pituitary NETs, and parathyroid tumors are at variance with the in vitro effects of menin, and it may be that menin interacts with proteins that inhibit apoptosis in these tissues or that menin does not have a significant in vivo role in apoptosis in these glands and their tumors.

Further investigation of apoptotic pathways in this model may therefore elucidate the different in vivo mechanisms involved.

Finally, use of long-term BrdU to label proliferating cells in vivo and thereby assess proliferation rates may help in evaluating the effectiveness of novel treatments aimed at reducing NET growth, such as monoclonal antibodies, gene therapy, and other genetic modifying agents.

This work was supported by the United Kingdom Medical Research Council Grants G, and G, to G. Thakker RV Multiple endocrine neoplasia type 1 MEN1.

Best Pract Res Clin Endocrinol Metab 24 : — Google Scholar. Newey PJ , Thakker RV Role of multiple endocrine neoplasia type 1 mutational analysis in clinical practice.

Endocr Pract 17 Suppl 3 : 8 — Calender A , Giraud S , Cougard P , Chanson P , Lenoir G , Murat A , Hamon P , Proye C Multiple endocrine neoplasia type 1 in France: clinical and genetic studies.

J Intern Med : — Trump D , Farren B , Wooding C , Pang JT , Besser GM , Buchanan KD , Edwards CR , Heath DA , Jackson CE , Jansen S , Lips K , Monson JP , O'Halloran D , Sampson J , Shalet SM , Wheeler MH , Zink A , Thakker RV Clinical studies of multiple endocrine neoplasia type 1 MEN1.

QJM 89 : — Lemmens I , Van de Ven WJ , Kas K , Zhang CX , Giraud S , Wautot V , Buisson N , De Witte K , Salandre J , Lenoir G , Pugeat M , Calender A , Parente F , Quincey D , Gaudray P , De Wit MJ , Lips CJ , Höppener JW , Khodaei S , Grant AL , Weber G , Kytölä S , Teh BT , Farnebo F , Thakker RV Identification of the multiple endocrine neoplasia type 1 MEN1 gene.

The European consortium on MEN1. Hum Mol Genet 6 : — Chandrasekharappa SC , Guru SC , Manickam P , Olufemi SE , Collins FS , Emmert-Buck MR , Debelenko LV , Zhuang Z , Lubensky IA , Liotta LA , Crabtree JS , Wang Y , Roe BA , Weisemann J , Boguski MS , Agarwal SK , Kester MB , Kim YS , Heppner C , Dong Q , Spiegel AM , Burns AL , Marx SJ Positional cloning of the gene for multiple endocrine neoplasia-type 1.

Science : — Schnepp RW , Chen YX , Wang H , Cash T , Silva A , Diehl JA , Brown E , Hua X Mutation of tumor suppressor gene Men1 acutely enhances proliferation of pancreatic islet cells.

Cancer Res 66 : — Mol Cell Endocrinol : 18 — Bazzi W , Renon M , Vercherat C , Hamze Z , Lacheretz-Bernigaud A , Wang H , Blanc M , Roche C , Calender A , Chayvialle JA , Scoazec JY , Cordier-Bussat M MEN1 missense mutations impair sensitization to apoptosis induced by wild-type menin in endocrine pancreatic tumor cells.

Gastroenterology : — Milne TA , Hughes CM , Lloyd R , Yang Z , Rozenblatt-Rosen O , Dou Y , Schnepp RW , Krankel C , Livolsi VA , Gibbs D , Hua X , Roeder RG , Meyerson M , Hess JL Menin and MLL cooperatively regulate expression of cyclin-dependent kinase inhibitors.

Proc Natl Acad Sci USA : — Eur J Cancer 43 : — La P , Yang Y , Karnik SK , Silva AC , Schnepp RW , Kim SK , Hua X Menin-mediated caspase 8 expression in suppressing multiple endocrine neoplasia type 1.

J Biol Chem : — Harding B , Lemos MC , Reed AA , Walls GV , Jeyabalan J , Bowl MR , Tateossian H , Sullivan N , Hough T , Fraser WD , Ansorge O , Cheeseman MT , Thakker RV Multiple endocrine neoplasia type 1 knockout mice develop parathyroid, pancreatic, pituitary and adrenal tumours with hypercalcaemia, hypophosphataemia and hypercorticosteronaemia.

Endocr Relat Cancer 16 : — Dolbeare F , Gratzner H , Pallavicini MG , Gray JW Flow cytometric measurement of total DNA content and incorporated bromodeoxyuridine.

Proc Natl Acad Sci USA 80 : — Diabetes 58 : — Diabetes 54 : — Bading JR , Shields AF Imaging of cell proliferation: status and prospects.

J Nucl Med 49 Suppl 2 : 64S — 80S. J Histochem Cytochem 35 : — Grigorieva IV , Mirczuk S , Gaynor KU , Nesbit MA , Grigorieva EF , Wei Q , Ali A , Fairclough RJ , Stacey JM , Stechman MJ , Mihai R , Kurek D , Fraser WD , Hough T , Condie BG , Manley N , Grosveld F , Thakker RV Gata3-deficient mice develop parathyroid abnormalities due to dysregulation of the parathyroid-specific transcription factor Gcm2.

J Clin Invest : — Gaffney EA The application of mathematical modelling to aspects of adjuvant chemotherapy scheduling. J Math Biol 48 : — Bender EA Introduction to mathematical modelling.

Mineola, New York : Dover Publications, Inc. Google Preview. Kidd M , Modlin IM Small intestinal neuroendocrine cell pathobiology: 'carcinoid' tumors.

Curr Opin Oncol 23 : 45 — J Surg Res : 68 — Sen A , Capitano ML , Spernyak JA , Schueckler JT , Thomas S , Singh AK , Evans SS , Hylander BL , Repasky EA Mild elevation of body temperature reduces tumor interstitial fluid pressure and hypoxia and enhances efficacy of radiotherapy in murine tumor models.

Cancer Res 71 : — Svejda B , Kidd M , Giovinazzo F , Eltawil K , Gustafsson BI , Pfragner R , Modlin IM The 5-HT 2B receptor plays a key regulatory role in both neuroendocrine tumor cell proliferation and the modulation of the fibroblast component of the neoplastic microenvironment.

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