Epigenetic origins of metabolic disease: The impact of the maternal condition to the offspring epigenome and later health consequences

D.J. Barker, C. Osmond, J. Golding, et al., Growth in utero, blood pressure in childhood and adult life, and mortality from cardiovascular disease, British Medical Journal 298 (1989) 564–567.

D.J. Barker, P.D. Winter, C. Osmond, et al., Weight in infancy and death from ischaemic heart disease, Lancet 2 (1989) 577–580.

H.O. Lithell, P.M. McKeigue, L. Berglund, et al., Relation of size at birth to non-insulin dependent diabetes and insulin concentrations in men aged 50–60 years, British Medical Journal 312 (1996) 406–410.

D.J. Barker, C.N. Hales, C.H. Fall, et al., Type 2 (non-insulin-dependent) diabetes mellitus, hypertension and hyperlipidaemia (syndrome X): relation to reduced fetal growth, Diabetologia 36 (1993) 62–67.

C.N. Hales, D.J. Barker, P.M. Clark, et al., Fetal and infant growth and impaired glucose tolerance at age 64, British Medical Journal 303 (1991) 1019–1022.

C.B. Jensen, H. Storgaard, S. Madsbad, et al., Altered skeletal muscle fiber composition and size precede whole-body insulin resistance in young men with low birth weight, The Journal of Clinical Endocrinology and Metabolism 92 (2007) 1530–1534.

J.G. Eriksson, C. Osmond, E. Kajantie, et al., Patterns of growth among children who later develop type 2 diabetes or its risk factors, Diabetologia 49 (2006) 2853–2858.

J.G. Eriksson, T. Forsén, J. Tuomilehto, et al., Effects of size at birth and childhood growth on the insulin resistance syndrome in elderly individuals, Diabetologia 45 (2002) 342–348.

C. Cooper, K. Walker-Bone, N. Arden, et al., Novel insights into the pathogenesis of osteoporosis: the role of intrauterine programming, Rheumatology 39 (2000) 1312–1315.

C. Cooper, C. Fall, P. Egger, et al., Growth in infancy and bone mass in later life, Annals of the Rheumatic Diseases 56 (1997) 17–21.

C.N. Hales, D.J. Barker, Type 2 (non-insulin-dependent) diabetes mellitus: the thrifty phenotype hypothesis, Diabetologia 35 (1992) 595–601.

P.D. Gluckman, W. Cutfield, P. Hofman, et al., The fetal, neonatal, and infant environments-the long-term consequences for disease risk, Early Human Development 81 (2005) 51–59.

P.D. Gluckman, M.A. Hanson, Maternal constraint of fetal growth and its consequences, Seminars in Fetal & Neonatal Medicine 9 (2004) 419–425.

P.D. Gluckman, M.A. Hanson, H.G. Spencer, Predictive adaptive responses and human evolution, Trends in Ecology & Evolution 20 (2005) 527–533.

P.D. Gluckman, M.A. Hanson, A.S. Beedle, et al., Predictive adaptive responses in perspective, Trends in Endocrinology and Metabolism 19 (2008) 109–110.

P.D. Gluckman, M.A. Hanson, The developmental origins of the metabolic syndrome, Trends in Endocrinology and Metabolism 15 (2004) 183–187.

M. Desai, C.N. Hales, Role of fetal and infant growth in programming metabolism in later life, Biological Reviews of the Cambridge Philosophical Society 72 (1997) 329–348.

P.M. Catalano, L. Presley, J. Minium, et al., Fetuses of obese mothers develop insulin resistance in utero, Diabetes Care 32 (2009) 1076–1080.

G. Mingrone, M. Manco, M.E.V. Mora, et al., Influence of maternal obesity on insulin sensitivity and secretion in offspring, Diabetes Care 31 (2008) 1872–1876.

D. Dabelea, E.J. Mayer-Davis, A.P. Lamichhane, et al., Association of intrauterine exposure to maternal diabetes and obesity with type 2 diabetes in youth: the SEARCH case-control study, Diabetes Care 31 (2008) 1422–1426.

J.C. King, Maternal obesity, metabolism, and pregnancy outcomes, Annual Review of Nutrition 26 (2006) 271–291.

A.J. Buckley, B. Keserü, J. Briody, et al., Altered body composition and metabolism in the male offspring of high fat-fed rats, Metabolism: Clinical and Experimental 54 (2005) 500–507.

J.C. Challier, S. Basu, T. Bintein, et al., Obesity in pregnancy stimulates macrophage accumulation and inflammation in the placenta, Placenta 29 (2008) 274–281.

J. Han, J. Xu, P.N. Epstein, et al., Long-term effect of maternal obesity on pancreatic beta cells of offspring: reduced beta cell adaptation to high glucose and high-fat diet challenges in adult female mouse offspring, Diabetologia 48 (2005) 1810–1818.

A.M. Samuelsson, P.A. Matthews, M. Argenton, et al., Diet-induced obesity in female mice leads to offspring hyperphagia, adiposity, hypertension, and insulin resistance: a novel murine model of developmental programming, Hypertension 51 (2008) 383–392.

J.F. Tong, X. Yan, M.J. Zhu, et al., Maternal obesity downregulates myogenesis and beta-catenin signaling in fetal skeletal muscle, American Journal of Physiology 296 (2009) E917–E924.

M.J. Zhu, B. Han, J. Tong, et al., AMP-activated protein kinase signalling pathways are down regulated and skeletal muscle development impaired in fetuses of obese, over-nourished sheep, The Journal of Physiology 586 (2008) 2651–2664.

M.J. Zhu, M. Du, P.W. Nathanielsz, et al., Maternal obesity up-regulates inflammatory signaling pathways and enhances cytokine expression in the mid-gestation sheep placenta, Placenta 31 (2010) 387–391.

J. Férézou-Viala, A.-F. Roy, C. Sérougne, et al., Long-term consequences of maternal high-fat feeding on hypothalamic leptin sensitivity and diet-induced obesity in the offspring, American Journal of Physiology 293 (2007) R1056–R1062.

B.M. Gregorio, V. Souza-Mello, J.J. Carvalho, et al., Maternal high-fat intake predisposes nonalcoholic fatty liver disease in C57BL/6 offspring, American Journal of Obstetrics and Gynecology 203 (2010), 495.e1-e8.

F. Guo, K.L. Jen, High-fat feeding during pregnancy and lactation affects offspring metabolism in rats, Physiology & Behavior 57 (1995) 681–686.

H.S. Kahn, K.M. Narayan, D.F. Williamson, et al., Relation of birth weight to lean and fat thigh tissue in young men, International Journal of Obesity and Related Metabolic Disorders 24 (2000) 667–672.

B.S. Muhlhausler, C.L. Adam, P.A. Findlay, et al., Increased maternal nutrition alters development of the appetite-regulating network in the brain, FASEB Journal 20 (2006) 1257–1259.

P.D. Taylor, J. McConnell, I.Y. Khan, et al., Impaired glucose homeostasis and mitochondrial abnormalities in offspring of rats fed a fat-rich diet in pregnancy, American Journal of Physiology 288 (2005) R134–R139.

C.L. White, M.N. Purpera, C.D. Morrison, Maternal obesity is necessary for programming effect of high-fat diet on offspring, American Journal of Physiology 296 (2009) R1464–R1472.

X. Yan, M.J. Zhu, W. Xu, et al., Up-regulation of Toll-like receptor 4/nuclear factor-kappaB signaling is associated with enhanced adipogenesis and insulin resistance in fetal skeletal muscle of obese sheep at late gestation, Endocrinology 151 (2010) 380–387.

L. Zhang, N.M. Long, S.M. Hein, et al., Maternal obesity in ewes results in reduced fetal pancreatic β-cell numbers in late gestation and decreased circulating insulin concentration at term, Domestic Animal Endocrinology 40 (2011) 30–39.

J.A. Oben, T. Patel, A. Mouralidarane, et al., Maternal obesity programmes offspring development of non-alcoholic fatty pancreas disease, Biochemical and Biophysical Research Communications 394 (2010) 24–28.

K. Hartil, P.M. Vuguin, M. Kruse, et al., Maternal substrate utilization programs the development of the metabolic syndrome in male mice exposed to high fat in utero, Pediatric Research 66 (2009) 368–373.

M. Srinivasan, S.D. Katewa, A. Palaniyappan, et al., Maternal high-fat diet consumption results in fetal malprogramming predisposing to the onset of metabolic syndrome-like phenotype in adulthood, American Journal of Physiology 291 (2006) E792–E799.

M.M. Elahi, F.R. Cagampang, D. Mukhtar, et al., Long-term maternal high-fat feeding from weaning through pregnancy and lactation predisposes offspring to hypertension, raised plasma lipids and fatty liver in mice, The British Journal of Nutrition 102 (2009) 514–519.

M.J. Zhu, Y. Ma, N.M. Long, et al., Maternal obesity markedly increases placental fatty acid transporter expression and fetal blood triglycerides at midgestation in the ewe, American Journal of Physiology 299 (2010) R1224–R1231.

S.E. Shoelson, L. Herrero, A. Naaz, Obesity, inflammation, and insulin resistance, Gastroenterology 132 (2007) 2169–2180.

G.M. Jonakait, The effects of maternal inflammation on neuronal development: possible mechanisms, International Journal of Developmental Neuroscience 25 (2007) 415–425.

E. Ardite, J.A. Barbera, J. Roca, et al., Glutathione depletion impairs myogenic differentiation of murine skeletal muscle C2C12 cells through sustained NF-kappaB activation, The American Journal of Pathology 165 (2004) 719–728.

H. Wang, E. Hertlein, N. Bakkar, et al., NF-kappaB regulation of YY1 inhibits skeletal myogenesis through transcriptional silencing of myofibrillar genes, Molecular and Cellular Biology 27 (2007) 4374–4387.

S.A. Bayol, B.H. Simbi, J.A. Bertrand, et al., Offspring from mothers fed a “junk food” diet in pregnancy and lactation exhibit exacerbated adiposity that is more pronounced in females, The Journal of Physiology 586 (2008) 3219–3230.

R.A. DeFronzo, E. Ferrannini, Y. Sato, et al., Synergistic interaction between exercise and insulin on peripheral glucose uptake, The Journal of Clinical Investigation 68 (1981) 1468–1474.

A.A. Sayer, H.E. Syddall, E.M. Dennison, et al., Grip strength and the metabolic syndrome: findings from the Hertfordshire Cohort Study, QJM: Monthly Journal of the Association of Physicians 100 (2007) 707–713.

D.I. Phillips, Relation of fetal growth to adult muscle mass and glucose tolerance, Diabetic Medicine 12 (1995) 686–690.

H. Ylihärsilä, E. Kajantie, C. Osmond, et al., Birth size, adult body composition and muscle strength in later life, International Journal of Obesity 31 (2007) 1392–1399.

D. Kuh, J. Bassey, R. Hardy, et al., Birth weight, childhood size, and muscle strength in adult life: evidence from a birth cohort study, American Journal of Epidemiology 156 (2002) 627–633.

C.B. Jensen, M.S. Martin-Gronert, H. Storgaard, et al., Altered PI3-kinase/Akt signalling in skeletal muscle of young men with low birth weight, PloS ONE 3 (2008) e3738.

S.E. Ozanne, C.B. Jensen, K.J. Tingey, et al., Decreased protein levels of key insulin signalling molecules in adipose tissue from young men with a low birthweight: potential link to increased risk of diabetes? Diabetologia 49 (2006) 2993–2999.

S.E. Ozanne, C.B. Jensen, K.J. Tingey, et al., Low birthweight is associated with specific changes in muscle insulin-signalling protein expression, Diabetologia 48 (2005) 547–552.

M.J. Zhu, S.P. Ford, W.J. Means, et al., Maternal nutrient restriction affects properties of skeletal muscle in offspring, The Journal of Physiology 575 (2006) 241–250.

P. Shelley, M.S. Martin-Gronert, A. Rowlerson, et al., Altered skeletal muscle insulin signaling and mitochondrial complex Ⅱ–Ⅲ linked activity in adult offspring of obese mice, American Journal of Physiology 297 (2009) R675–R681.

R. Bergeron, R.R. Russell, L.H. Young, et al., Effect of AMPK activation on muscle glucose metabolism in conscious rats, The American Journal of Physiology 276 (1999) E938–E944.

T. Vaissière, C. Sawan, Z. Herceg, Epigenetic interplay between histone modifications and DNAmethylation in gene silencing, Mutation Research 659 (659) 40–48.

A.J. Bannister, T. Kouzarides, Reversing histone methylation, Nature 436 (2005) 1103–1106.

D. Sproul, N. Gilbert, W.A. Bickmore, The role of chromatin structure in regulating the expression of clustered genes, Nature Reviews: Genetics 6 (2005) 775–781.

D. Schübeler, M.C. Lorincz, D.M. Cimbora, et al., Genomic targeting of methylated DNA: influence of methylation on transcription, replication, chromatin structure, and histone acetylation, Molecular and Cellular Biology 20 (2000) 9103–9112.

A.P. Wolffe, M.A. Matzke, Epigenetics: regulation through repression, Science 286 (1999) 481–486.

C. Sawan, T. Vaissière, R. Murr, et al., Epigenetic drivers and genetic passengers on the road to cancer, Mutation Research 642 (2008) 1–13.

S. Dedeurwaerder, F. Fuks, DNA methylation markers for breast cancer prognosis: unmasking the immune component, Oncoimmunology 1 (2012) 962–964.

D. Bernard, J. Gil, P. Dumont, et al., The methyl-CpG-binding protein MECP2 is required for prostate cancer cell growth, Oncogene 25 (2006) 1358–1366.

O. Babenko, I. Kovalchuk, G. Metz, Epigenetic programming of neurodegenerative diseases by an adverse environment, Brain Research 1444 (2012) 96–111.

S. Marques, T.F. Outeiro, Epigenetics in Parkinson’s and Alzheimer’s diseases, Sub-cellular Biochemistry 61 (2012) 507–525.

M. Volkmar, S. Dedeurwaerder, D.A. Cunha, et al., DNA methylation profiling identifies epigenetic dysregulation in pancreatic islets from type 2 diabetic patients, The EMBO Journal 31 (2012) 1405–1426.

C. Brøns, S. Jacobsen, E. Nilsson, et al., Deoxyribonucleic acid methylation and gene expression of PPARGC1A in human muscle is influenced by high-fat overfeeding in a birth-weight-dependent manner, The Journal of Clinical Endocrinology and Metabolism 95 (2010) 3048–3056.

C. Ling, S. Del Guerra, R. Lupi, et al., Epigenetic regulation of PPARGC1A in human type 2 diabetic islets and effect on insulin secretion, Diabetologia 51 (2008) 615–622.

R. Barrès, M.E. Osler, J. Yan, et al., Non-CpG methylation of the PGC-1alpha promoter through DNMT3B controls mitochondrial density, Cell Metabolism 10 (2009) 189–198.

R. Benetti, S. Gonzalo, I. Jaco, et al., A mammalian microRNA cluster controls DNA methylation and telomere recombination via Rbl2-dependent regulation of DNA methyltransferases, Nature Structural & Molecular Biology 15 (2008) 268–279.

L. Sinkkonen, T. Hugenschmidt, P. Berninger, et al., MicroRNAs control de novo DNA methylation through regulation of transcriptional repressors in mouse embryonic stem cells, Nature Structural & Molecular Biology 15 (2008) 259–267.

W. Reik, Stability and flexibility of epigenetic gene regulation in mammalian development, Nature 447 (2007) 425–432.

N. Bhutani, D.M. Burns, H.M. Blau, DNA demethylation dynamics, Cell 146 (2011) 866–872.

S.E. Pinney, L.J. Jaeckle Santos, Y. Han, et al., Exendin-4 increases histone acetylase activity and reverses epigenetic modifications that silence Pdx1 in the intrauterine growth retarded rat, Diabetologia 54 (2011) 2606–2614.

J.H. Park, D.A. Stoffers, R.D. Nicholls, et al., Development of type 2 diabetes following intrauterine growth retardation in rats is associated with progressive epigenetic silencing of Pdx1, The Journal of Clinical Investigation 118 (2008) 2316–2324.

R.F. Thompson, F.H. Einstein, Epigenetic basis for fetal origins of age-related disease, Journal of Women’s Health 19 (2010) 581–587.

J.L. Slater-Jefferies, K.A. Lillycrop, P.A. Townsend, et al., Feeding a protein-restricted diet during pregnancy induces altered epigenetic regulation of peroxisomal proliferator-activated receptor-α in the heart of the offspring, Journal of Developmental Origins of Health and Disease 2 (2011) 250–255.

K.A. Lillycrop, J.L. Slater-Jefferies, M.A. Hanson, et al., Induction of altered epigenetic regulation of the hepatic glucocorticoid receptor in the offspring of rats fed a protein-restricted diet during pregnancy suggests that reduced DNA methyltransferase-1 expression is involved in impaired DNA methylation and changes in histone modifications, The British Journal of Nutrition 97 (2007) 1064–1073.

S.E. Pinney, R.A. Simmons, Epigenetic mechanisms in the development of type 2 diabetes, Trends in Endocrinology and Metabolism 21 (2010) 223–229.

F. Fuks, DNA methylation and histone modifications: teaming up to silence genes, Current Opinion in Genetics & Development 15 (2005) 490–495.

F. Fuks, P.J. Hurd, D. Wolf, et al., The methyl-CpG-binding protein MeCP2 links DNA methylation to histone methylation, The Journal of Biological Chemistry 278 (2003) 4035–4040.

A. Bird, DNA methylation patterns and epigenetic memory, Genes & Development 16 (2002) 6–21.

J. Trasler, L. Deng, S. Melnyk, et al., Impact of Dnmt1 deficiency, with and without low folate diets, on tumor numbers and DNA methylation in Min mice, Carcinogenesis 24 (2003) 39–45.

M. Fabbri, R. Garzon, A. Cimmino, et al., MicroRNA-29 family reverts aberrant methylation in lung cancer by targeting DNA methyltransferases 3A and 3B, Proceedings of the National Academy of Sciences of the United States of America 104 (2007) 15805–15810.

J.F. Glickman, J.G. Pavlovich, N.O. Reich, Peptide mapping of the murine DNA methyltransferase reveals a major phosphorylation site and the start of translation, The Journal of Biological Chemistry 272 (1997) 17851–17857.

R. Goyal, P. Rathert, H. Laser, et al., Phosphorylation of serine-515 activates the Mammalian maintenance methyltransferase Dnmt1, Epigenetics 2 (2007) 155–160.

H. Denis, M.N. Ndlovu, F. Fuks, Regulation of mammalian DNA methyltransferases: a route to new mechanisms, EMBO Reports 12 (2011) 647–656.

O. Bogdanović, G.J.C. Veenstra, DNA methylation and methyl-CpG binding proteins: developmental requirements and function, Chromosoma 118 (2009) 549–565.

D.M. Woodcock, P.J. Crowther, W.P. Diver, The majority of methylated deoxycytidines in human DNA are not in the CpG dinucleotide, Biochemical and Biophysical Research Communications 145 (1987) 888–894.

M.J. Ziller, F. Müller, J. Liao, et al., Genomic distribution and intersample variation of non-CpG methylation across human cell types, PLoS Genetics 7 (2011) e1002389.

B.H. Ramsahoye, D. Biniszkiewicz, F. Lyko, et al., Non-CpG methylation is prevalent in embryonic stem cells and may be mediated by DNA methyltransferase 3a, Proceedings of the National Academy of Sciences of the United States of America 97 (2000) 5237–5242.

S. Sookoian, M.S. Rosselli, C. Gemma, et al., Epigenetic regulation of insulin resistance in nonalcoholic fatty liver disease: impact of liver methylation of the peroxisome proliferator-activated receptor γ coactivator 1α promoter, Hepatology 52 (2010) 1992–2000.

H.D. Morgan, H.G. Sutherland, D.I. Martin, et al., Epigenetic inheritance at the agouti locus in the mouse, Nature genetics 23 (1999) 314–318.

G.L. Wolff, R.L. Kodell, S.R. Moore, et al., Maternal epigenetics and methyl supplements affect agouti gene expression in Avy/a mice, FASEB Journal 12 (1998) 949–957.

J.E. Cropley, C.M. Suter, K.B. Beckman, et al., Germ-line epigenetic modification of the murine A vy allele by nutritional supplementation, Proceedings of the National Academy of Sciences of the United States of America 103 (2006) 17308–17312.

G.C. Burdge, J. Slater-Jefferies, C. Torrens, et al., Dietary protein restriction of pregnant rats in the F0 generation induces altered methylation of hepatic gene promoters in the adult male offspring in the F1 and F2 generations, The British Journal of Nutrition 97 (2007) 435–439.

G.C. Burdge, S.P. Hoile, T. Uller, et al., Progressive, transgenerational changes in offspring phenotype and epigenotype following nutritional transition, PloS ONE 6 (2011) e28282.

C. Gemma, S. Sookoian, J. Alvariñas, et al., Maternal pregestational BMI is associated with methylation of the PPARGC1A promoter in newborns, Obesity 17 (2009) 1032–1039.

T. Forsén, J. Eriksson, J. Tuomilehto, et al., The fetal and childhood growth of persons who develop type 2 diabetes, Annals of Internal Medicine 133 (2000) 176–182.

P. Völkel, P.O. Angrand, The control of histone lysine methylation in epigenetic regulation, Biochimie 89 (2007) 1–20.

J.C. Rice, C.D. Allis, Histone methylation versus histone acetylation: new insights into epigenetic regulation, Current Opinion in Cell Biology 13 (2001) 263–273.

Q. Fu, R.A. McKnight, X. Yu, et al., Uteroplacental insufficiency induces site-specific changes in histone H3 covalent modifications and affects DNA-histone H3 positioning in day 0 IUGR rat liver, Physiological Genomics 20 (2004) 108–116.

K.M. Aagaard-Tillery, K. Grove, J. Bishop, et al., Developmental origins of disease and determinants of chromatin structure: maternal diet modifies the primate fetal epigenome, Journal of Molecular Endocrinology 41 (2008) 91–102.

R.S. Strakovsky, X. Zhang, D. Zhou, et al., Gestational high fat diet programs hepatic phosphoenolpyruvate carboxykinase gene expression and histone modification in neonatal offspring rats, The Journal of Physiology 589 (2011) 2707–2717.

N. Raychaudhuri, S. Raychaudhuri, M. Thamotharan, et al., Histone code modifications repress glucose transporter 4 expression in the intrauterine growth-restricted offspring, The Journal of Biological Chemistry 283 (2008) 13611–13626.

S. Zheng, M. Rollet, Y.X. Pan, Maternal protein restriction during pregnancy induces CCAAT/enhancer-binding protein (C/EBPβ) expression through the regulation of histone modification at its promoter region in female offspring rat skeletal muscle, Epigenetics 6 (2011) 161–170.

J. Zhang, F. Zhang, X. Didelot, et al., Maternal high fat diet during pregnancy and lactation alters hepatic expression of insulin like growth factor-2 and key microRNAs in the adult offspring, BMC Genomics 10 (2009) 478.

A.L. Siebel, A. Mibus, M.J. De Blasio, et al., Improved lactational nutrition and postnatal growth ameliorates impairment of glucose tolerance by uteroplacental insufficiency in male rat offspring, Endocrinology 149 (2008) 3067–3076.

K.S. Park, S.K. Kim, M.S. Kim, et al., Fetal and early postnatal protein malnutrition cause long-term changes in rat liver and muscle mitochondria, The Journal of Nutrition 133 (2003) 3085–3090.

R.C. Laker, L.A. Gallo, M.E. Wlodek, et al., Short-term exercise training early in life restores deficits in pancreatic β-cell mass associated with growth restriction in adult male rats, American Journal of Physiology 301 (2011) E931–E940.

R.C. Laker, M.E. Wlodek, G.D. Wadley, et al., Exercise early in life in rats born small does not normalize reductions in skeletal muscle PGC-1α in adulthood, American journal of physiology, Endocrinology and Metabolism 302 (2012) E1221–E1230.

M. Garg, M. Thamotharan, S.A. Oak, et al., Early exercise regimen improves insulin sensitivity in the intrauterine growth-restricted adult female rat offspring, American Journal of Physiology 296 (2009) E272–E281.

J.L. Miles, K. Huber, N.M. Thompson, et al., Moderate daily exercise activates metabolic flexibility to prevent prenatally induced obesity, Endocrinology 150 (2009) 179–186.

K. Huber, J.L. Miles, A.M. Norman, et al., Prenatally induced changes in muscle structure and metabolic function facilitate exercise-induced obesity prevention, Endocrinology 150 (2009) 4135–4144.

J.G. Eriksson, H. Ylihärsilä, T. Forsén, et al., Exercise protects against glucose intolerance in individuals with a small body size at birth, Preventive Medicine 39 (2004) 164–167.

B.A. Irving, K.R. Short, K.S. Nair, et al., Nine days of intensive exercise training improves mitochondrial function but not insulin action in adult offspring of mothers with type 2 diabetes, The Journal of Clinical Endocrinology and Metabolism 96 (2011) E1137–E1141.

M.E. Wlodek, A. Mibus, A. Tan, et al., Normal lactational environment restores nephron endowment and prevents hypertension after placental restriction in the rat, Journal of the American Society of Nephrology 18 (2007) 1688–1696.

L.G. Carter, K.N. Lewis, D.C. Wilkerson, et al., Perinatal exercise improves glucose homeostasis in adult offspring, American Journal of Physiology, Endocrinology and Metabolism 303 (2012) E1061–E1068.

L.G. Carter, N.R. Qi, R. De Cabo, et al., Maternal exercise improves insulin sensitivity in mature rat offspring, Medicine and Science in Sports and Exercise (2012), PMID 23247711.

S.P. Hoile, N.A. Irvine, C.J. Kelsall, et al., Maternal fat intake in rats alters 20:4n − 6 and 22:6n − 3 status and the epigenetic regulation of Fads2 in offspring liver, The Journal of Nutritional Biochemistry (2012), http://dx.doi.org/10.1016/j.jnutbio.2012.09.005.