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Richard Weindruch

Richard Weindruch

Ph.D., University of California-Los Angeles
Professor Emeritus, Department of Medicine- Geriatrics

Influences of Calorie Intake on Aging

It has been known for 65 years that caloric restriction (CR), if properly executed without malnutrition, will greatly extend the maximum life span of mice and rats and decrease the incidence of cancer and other late-life diseases (for a review of CR, see Weindruch and Walford, 1988). Although regarded more as a curiosity for much of that time, interest in CR-induced phenomena grew substantially with increased research on aging processes since the 1970s and continues to this day.

Most of the CR studies carried out in my laboratory have been in mice from long-lived strains that develop cancer (lymphoma, hepatoma) late in life. Mice on CR eat up to 50% fewer calories than controls while consuming the same amounts of proteins, vitamins and minerals. In one of our studies, CR increased the average life span from 32 months to 45 months, and increased by 34% the maximum (last decile) life span from 40 months to 53 months. Overall tumor incidence was 38% in the CR group vs. 78% in the control group. In addition to living longer, we found that mice on CR stay "younger longer" than controls as judged by such age-sensitive biologic parameters as immune responses, eye lens proteins, liver enzyme activities, learning and behavior.

Two main questions now face this field of research and serve to focus the efforts of this laboratory. These are: 1) What are the mechanisms underlying the retardation of aging and disease processes by CR in rodents?, and 2) Does CR also act to retard aging and diseases in nonhuman and human primates?

Our present work in rodents involves testing the possibility that CR acts by reducing oxidative stress in postmitotic tissues. In 1998, a collaboration began with Dr. Tomas Prolla using oligonucleotide microarrays to conduct the first gene expression profiling studies in gerontology. Data from these studies strengthen the idea that CR may slow aging by reducing free radical damage to tissues. Whether this mechanism is operative in opposing late-life cancers is worthy of expanded exploration.

My laboratory is also exploring the efficacy of CR initiated in middle age (or later) to retard aging processes in nonhuman primates. These studies are being done in collaboration with Drs. Kemnitz, Schoeller and others who are exploring age-sensitive parameters in adult monkeys subjected to CR of 30%. The feasibility of human studies is under active consideration.

Representative Publications
Barger, J. L., Anderson, R. M., Newton, M. A., da Silva, C., Vann, J. A., Pugh, T. D., Someya, S., Prolla, T. A., & Weindruch, R (2015). A conserved transcriptional signature of delayed aging and reduced disease vulnerability is partially mediated by SIRT3. PLoS One, 10(4), e0120738.
View publication via DOI: DOI:10.1371/journal.pone.0120738

Colman, R.J., Beasley, T.M., Kemnitz, J.W., Johnson, S.C., Weindruch, R., & Anderson, R.M. (2014). Caloric restriction increases age-related and all cause survival in rhesus monkeys. Nature Communications 5, 3557.
View publication via DOI: DOI:10.1038/ncomms4557

Sridharan, A., Bendlin, B. B., Gallagher, C. L., Oh, J. M., Willette, A. A., Alexander, A. L., Kemnitz, J. W., Colman, R. J., Weindruch, R. H., & Johnson, S. C. (2014). Effect of age and calorie restriction on corpus callosal integrity in rhesus macaques: A fiber tractography study. Neuroscience Letters, 569, 38-42.
View publication via DOI: DOI:10.1016/j.neulet.2014.03.047

Sridharan, A., Pehar, M., Salamat, M. S., Pugh, T. D., Bendlin, B. B., Willette, A. A., Anderson, R. M., Kemnitz, J. W., . . . Weindruch, R., Puglielli, L., & Johnson, S. C. (2013). Calorie restriction attenuates astrogliosis but not amyloid plaque load in aged rhesus macaques: A preliminary quantitative imaging study. Brain Research, 1508, 1-8.
View publication via DOI: DOI:10.1016/j.brainres.2013.02.046

Yamada, Y., Colman, R. J., Kemnitz, J. W., Baum, S. T., Anderson, R. M., Weindruch, R., & Schoeller, D. A. (2013). Long-term calorie restriction decreases metabolic cost of movement and prevents decrease of physical activity during aging in rhesus monkeys. Experimental Gerontology, 48(11), 1226-1235.
View publication via DOI: DOI:10.1016/j.exger.2013.08.002

Pugh, T. D., Conklin, M. W., Evans, T. D., Polewski, M. A., Barbian, H. J., Pass, R., . . . Weindruch, R., . . . Anderson, R. M. (2013). A shift in energy metabolism anticipates the onset of sarcopenia in rhesus monkeys. Aging Cell, 12(4), 672-681.
View publication via DOI: DOI:10.1111/acel.12091

Collino, S., Martin, F.-P., Montoliu, I., Barger, J., DaSilva, L., Prolla, T., Weindruch, R., Kochhar, S. (2013). Transcriptomics and metabonomics identify essential metabolic signatures in caloric restriction regulation across multiple mouse strains. Metabolites, 3(4), 881-911.
View publication via DOI: DOI:10.3390/metabo3040881

Willette, A. A., Coe, C. L., Birdsill, A. C., Bendlin, B. B., Colman, R. J., Alexander, A. L., ... Weindruch, R. H., & Johnson, S. C. (2013). Interleukin-8 and interleukin-10, brain volume and microstructure, and the influence of calorie restriction in old rhesus macaques. Age, 35(6), 2215-2227.
View publication via DOI: DOI:10.1007/s11357-013-9518-y

Willette, A.A., Bendlin, B.B., Colman, R.J., Kastman, E.K., Field, A.S., Alexander, A.L., ... Kemnitz, J.W., Weindruch, R.H., & Johnson, S. (2012). Calorie restriction reduces the influence of glucoregulator dysfunction on regional brain volume in aged rhesus monkeys. Diabetes, 61(5), 1036-1042.
View publication via DOI: DOI:10.2337/db11-1187

Willette, A.A., Gallagher, C., Bendlin, B.B., McLaren, D.G., Kastman, E.K., Canu, E., ... Weindruch, R.H., Coe, C.L., & Johnson, S.C. (2012). Homocysteine, neural atrophy, and the effect of caloric restriction in rhesus monkeys. Neurobiology of Aging, 33(4), 670-680.
View publication via DOI: DOI:10.1016/j.neurobiolaging.2010.06.003

Anderson, R. M., & Weindruch, R. (2012). The caloric restriction paradigm: Implications for healthy human aging. Am J Hum Biol, 24(2), 101-106.
View publication via DOI: DOI:10.1002/ajhb.22243

Colman, R. J., Beasley, T. M., Allison, D. B., & Weindruch, R. (2012). Skeletal effects of long-term caloric restriction in rhesus monkeys. Age, 34(5), 1133-1143.
View publication via DOI: DOI:10.1007/s11357-011-9354-x

Csiszar, A., Sosnowska, D., Tucsek, Z., Gautam, T., Toth, P., Losonczy, G., . . . Weindruch, R., Anderson, R. M., & Sonntag, W. E.Ungvari, Z. (2012). Circulating factors induced by caloric restriction in the nonhuman primate macaca mulatta activate angiogenic processes in endothelial cells. Journals of Gerontology. Series A, Biological Sciences and Medical Sciences, 68(3), 235-249.
View publication via DOI: DOI:10.1093/gerona/gls158

McKiernan, S. H., Colman, R. J., Aiken, E., Evans, T. D., Beasley, T. M., Aiken, J. M., . . . Weindruch, R., & Anderson, R. M. (2012). Cellular adaptation contributes to calorie restriction-induced preservation of skeletal muscle in aged rhesus monkeys. Experimental Gerontology, 47(3), 229-236.
View publication via DOI: DOI:10.1016/j.exger.2011.12.009

Sridharan, A., Willette, A. A., Bendlin, B. B., Alexander, A. L., Coe, C. L., Voytko, M. L., Coleman, R. J., Kemnitz, J. W., Weindruch, R. H., & Johnson, S. C. (2012). Brain volumetric and microstructural correlates of executive and motor performance in aged rhesus monkeys. Front Aging Neurosci, 4, 31.
View publication via DOI: DOI:10.3389/fnagi.2012.00031

Willette, A. A., Coe, C. L., Colman, R. J., Bendlin, B. B., Kastman, E. K., Field, A. S., ... Weindruch, R. H., & Johnson, S. C. (2012). Calorie restriction reduces psychological stress reactivity and its association with brain volume and microstructure in aged rhesus monkeys. Psychoneuroendocrinology, 37(7), 903-916.
View publication via DOI: DOI:10.1016/j.psyneuen.2011.10.006

Bendlin, B. B., Canu, E., Willette, A., Kastman, E.K., McLaren, D.G., Kosmatka, K.J., ... Coe, C.L., Weindruch, R.H., Alexander, A.L., & Johnson, S. C. (2011). Effects of aging and calorie restriction on white matter in rhesus macaques. Neurobiology of Aging, 32(12), 2319.e1-2319.e11.
View publication via DOI: DOI:10.1016/j.neurobiolaging.2010.04.008

McKiernan, S. H., Colman, R. J., Lopez, M., Beasley, T. M., Aiken, J. M., Anderson, R. M., & Weindruch, R. (2011). Caloric restriction delays aging-induced cellular phenotypes in rhesus monkey skeletal muscle. Experimental Gerontology, 46(1), 23-29.
View publication via DOI: DOI:10.1016/j.exger.2010.09.011

Robertson, H. T., Smith, D. L., Pajewski, N. M., Weindruch, R. H., Garland, T., Jr., Argyropoulos, G., . . . Allison, D. B. (2011). Can rodent longevity studies be both short and powerful? Journals of Gerontology. Series A, Biological Sciences and Medical Sciences, 66(3), 279-286.
View publication via DOI: DOI:10.1093/gerona/glq190

Fowler, C.G., Chiasson, K.B., Leslie, T.H., Thomas, D., Beasley, T.M., Kemnitz, J.W., & Weindruch, R. (2010). Auditory function in rhesus monkeys: Effects of aging and caloric restriction in the Wisconsin monkeys five years later. Hearing Research, 261(1-2), 75-81.
Click here to download this publication.
View publication via DOI: DOI:10.1016/j.heares.2010.01.006

Anderson, R.M., & Weindruch, R. (2010). Metabolic reprogramming, caloric restriction and aging. Trends in Endocrinology and Metabolism, 21(3), 134-141.
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View publication via DOI: DOI:10.1016/j.tem.2009.11.005

Someya, S., Tanokura, M., Weindruch, R., Prolla, T.A., & Yamasoba, T. (2010). Effects of caloric restriction on age-related hearing loss in rodents and rhesus monkeys. Current Aging Science, 3(1), 20-25.
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Willette, A.A., Bendlin, B.B., McLaren, D.G., Canu, E., Kastman, E.K., Kosmatka, K.J., ... Weindruch, R.H., Coe, C.L., & Johnson, S.C. (2010). Age-related changes in neural volume and microstructure associated with interleukin-6 are ameliorated by a calorie-restricted diet in old rhesus monkeys. NeuroImage, 51(3), 987-994.
View publication via DOI: DOI:10.1016/j.neuroimage.2010.03.015

Kastman, E.K., Willette, A.A., Coe, C.L., Bendlin, B.B., Kosmatka, K.J., McLaren, D.G., ... Weindruch, R.H., & Johnson, S. C. (2010). A calorie-restricted diet decreases brain iron accumulation and preserves motor performance in old rhesus monkeys. The Journal of Neuroscience, 32(34), 11897-11904.
View publication via DOI: DOI:10.1523/JNEUROSCI.2553-12.2012

Colman, R.J., Anderson, R.M., Johnson, S.C., Kastman, E.K., Kosmatka, K.J., T. Beasley, T.M., Allison, D.B., Cruzen, C., Simmons, H.A., Kemnitz, J.W., & Weindruch, R. (2009). Caloric restriction delays disease onset and mortality in rhesus monkeys. Science, 325, 201-204.
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View publication via DOI: DOI:10.1126/science.1173635

Someya, S., Xu, J., Kondo, K., Ding, D., Salvi, R.J., Yamasoba, T., Rabinovitch, P.S., Weindruch, R., Leeuwenburgh, C., Tanokura, M., & Prolla, T.A. (2009). Age-related hearing loss in C57BL/6J mice is mediated by Bak-dependent mitochondrial apoptosis. Proceedings of the National Academy of Sciences, 106(46), 19432-19437.
View publication via DOI: DOI:10.1073/pnas.0908786106

Park, S.K., Kim, K., Page, G.P., Allison, D.B., Weindruch, R., & Prolla, T.A. (2009). Gene expression profiling of aging in multiple mouse strains: Identification of aging biomarkers and impact of dietary antioxidants. Aging Cell, 8(4), 484-495.
View publication via DOI: DOI:10.1111/j.1474-9726.2009.00496.x

Hagopian, K., Ramsey, J.J., & Weindruch, R. (2009). Caloric restriction counteracts age-related changes in the activities of sorbitol metabolizing enzymes from mouse liver. Biogerontology, 10(4), 471-479.
View publication via DOI: DOI:10.1007/s10522-008-9191-1

McKiernan, S.H., Colman, R., Lopez, M., Beasley, T.M., Weindruch, R., & Aiken, J.M. (2009). Longitudinal analysis of early stage sarcopenia in aging rhesus monkeys. Experimental Gerontology, 44(3), 170-176.
View publication via DOI: DOI:10.1016/j.exger.2008.09.014

Anderson, R.M., Shanmuganayagam, D., & Weindruch, R. (2009). Caloric restriction and aging: Studies in mice and monkeys. Toxicologic Pathology, 37(1), 47-51.
View publication via DOI: DOI:10.1177/0192623308329476

Li, H., Costantini, C., Scrable, H., Weindruch, R., & Puglielli, L. (2008). Egr-1 and Hipk2 are required for the TrkA to p75NTR switch that occurs downstream of IGF1-R. Neurobiology of Aging, 30(12), 2010-2020.
View publication via DOI: DOI:10.1016/j.neurobiolaging.2008.02.015

Anderson, R.M., Barger, J.L., Edwards, M.G., Braun, K.H., O'Connor, C.E., Prolla, T.A., & Weindruch, R. (2008). Dynamic regulation of PGC-1alpha localization and turnover implicates mitochondrial adaptation in longevity and the stress response. Aging Cell, 7, 10.
View publication via DOI: DOI:10.1111/j.1474-9726.2007.00357.x

Barger, J.L., Kayo, T., Pugh, T.D., Prolla, T.A., & Weindruch, R. (2008). Short-term consumption of a resveratrol-containing nutraceutical mixture mimics gene expression of long-term caloric restriction in mouse heart. Experimental Gerontology, 43(9), 859-86.
View publication via DOI: DOI:10.1016/j.exger.2008.06.013

Fu, V.X., Dobosy, J.R., Desotelle, J.A., Almassi, N., Ewald, J.A., Srinivasan, R., Berres, M., Svaren, J., Weindruch, R., & Jarrard, D.F. (2008). Aging and cancer-related loss of insulin-like growth factor 2 imprinting in the mouse and human prostate. Cancer Research, 68(16), 6797-680.
View publication via DOI: DOI:10.1158/0008-5472.CAN-08-1714

Dobosy, J.R., Fu, V.X., Desotelle, J.A., Srinivasan, R., Kenowski, M.L., Almassi, N., Weindruch, R., Svaren, J., & Jarrard, D.F. (2008). A methyl-deficient diet modifies histone methylation and alters Igf2 and H19 repression in the prostate. Prostate, 68(11), 1187-119.
View publication via DOI: DOI:10.1002/pros.20782

Colman, R.J., Beasley, T.M., Allison, D.B., & Weindruch, R. (2008). Attenuation of sarcopenia by dietary restriction in rhesus monkeys. Journals of Gerontology Series A- Biological Sciences and Medical Sciences, 63(6), 556-559.

Hagopian, K., Ramsey, J.J., & Weindruch, R. (2008). Enzymes of glycerol and glyceraldehyde metabolism in mouse liver: effects of caloric restriction and age on activities. Bioscience Reports, 28(2), 107-11.
View publication via DOI: DOI:10.1042/BSR20080015

Someya, S., Yamasoba, T., Kujoth, G.C., Pugh, T.D., Weindruch, R., Tanokura, M., & Prolla, T.A. (2008). The role of mtDNA mutations in the pathogenesis of age-related hearing loss in mice carrying a mutator DNA polymerase gamma. Neurobiology of Aging, 29(7), 1080-109.
View publication via DOI: DOI:10.1016/j.neurobiolaging.2007.01.014

Park, S.K., Page, G.P., Kim, K., Allison, D.B., Meydani, M., Weindruch, R., & Prolla, T.A. (2008). Alpha- and gamma-tocopherol prevent age-related transcriptional alterations in the heart and brain of mice. Journal of Nutrition, 138(6), 1010-1018.

Barger, J.L., Kayo, T., Vann, J.M., Arias, E.B., Wang, J., Hacker, T.A., Wang, Y., Raederstorff, D., Morrow, J.D., Leeuwenburgh, C., Allison, D.B., Saupe, K.W., Cartee, G.D., Weindruch, R., & Prolla, T.A. (2008). A low dose of dietary resveratrol partially mimics caloric restriction and retards aging parameters in mice. PLoS ONE, 3(6), e2264.
View publication via DOI: DOI:10.1371/journal.pone.0002264

Fowler, C.G., Chiasson, K.B., Hart, D.B., Beasley, T.M., Kemnitz, J., & Weindruch, R. (2008). Tympanometry in rhesus monkeys: Effects of aging and caloric restriction. Int. J. Audiol., 47(4), 209-14.
View publication via DOI: DOI:10.1080/14992020701851882

Fowler, C.G., Chaisson, K.B., Brown Hart, D., Kemnitz J., & Weindruch, R. (2007). Tympanometric findings in rhesus monkeys: Effects of aging and caloric restriction. International Journal of Audiology, 47(4), 209-214.
View publication via DOI: DOI:10.1080/14992020701851882

Edwards, M.G., Anderson, R.M., Yuan, M., Kendziorski, C., Weindruch, R., & Prolla, T.A. (2007). Gene expression profiling of aging reveals activation of a p53-mediated transcriptional program. BMC Genomics, 8, 80.
View publication via DOI: DOI:10.1186/1471-2164-8-80

Anderson, R.M., & Weindruch, R. (2007). Metabolic reprogramming in calorie restriction. In C.V. Mobbs, K. Yen, P.R. Hof., & A.G. Karger (Eds.), Mechanisms of calorie restriction in aging and disease (Interdisciplinary Topics in Gerontology) (Vol. 35, pp. 18-38). Switzerland.

Raman, A., Ramsey, J.J., Kemnitz, J.W., Baum, S.T., Newton, W., Coleman, R.J., Weindruch, R., Beasley, M.T., & Schoeller, D.A. (2007). Influences of calorie restriction and age on energy expenditure in the rhesus monkeys. Am. J. Physiol., 292, E101-6.
View publication via DOI: DOI:10.​1152/​ajpendo.​00127.​2006

Raman, A., Baum, S.T., Colman, R.J., Kemnitz, J.W., Weindruch, R., & Schoeller, D.A. (2007). Metabolizable energy intake during long-term calorie restriction in rhesus monkeys. Exp. Gerontol., 42(10), 988-94.
View publication via DOI: DOI:10.1016/j.exger.2007.05.008

Someya, S., Yamasoba, T., Weindruch, R., Prolla, T.A., & Tanokura, M. (2007). Caloric restriction suppresses apoptotic cell death in the mammalian cochlea and leads to prevention of presbycusis. Neurobiology of Aging, 28, 1613-1622.
View publication via DOI: DOI:10.1016/j.neurobiolaging.2006.06.024

Yamasoba, T., Someya, S., Yamada, C., Weindruch, R., Prolla, TA., & Tanokura, M. (2007). Role of mitochondrial dysfunction and mitochondrial DNA mutations in age-related hearing loss. Hearing Research, 26, 185-193.
View publication via DOI: DOI:10.1016/j.heares.2006.06.004

Anderson, R.M., & Weindruch, R. (2006). Calorie Restriction: Progress during mid-2005 to mid-2006. Exp. Gerontology, 41, 1247-9.
View publication via DOI: DOI:10.1016/j.exger.2006.10.019

Costantini, C., Weindruch, R., Della Valle, G., & Puglielli, L. (2005). A TrkA to p75NTR molecular switch activates amyloid beta-peptide generation during aging. Biochemical Journal, 391, 59-67.
View publication via DOI: DOI:10.1042/BJ20050700

Raman, A., Colman, R.J., Cheng, Y., Kemnitz, J.W., Baum, S.T., Weindruch, R., & Schoeller, D.A. (2005). Reference body composition in adult rhesus monkeys: Glucoregulatory and anthropometric indices. J. Gerontol. A. Biol. Sci. Med. Sci., 60(12), 1518-24.
View publication via DOI: DOI:10.1093/gerona/60.12.1518

Blanc, S., Colman, R., Kemnitz, J., Weindruch, R., Baum, S., Ramsey, J., & Schoeller, D. (2005). Assessment of nutritional status in rhesus monkeys: comparison of dual-energy X-ray absorptiometry and stable isotope dilution. J. Med. Primatol., 34(3), 130-8.
View publication via DOI: DOI:10.1111/j.1600-0684.2005.00106.x

Kujoth, G.C., Hiona, A., Pugh, T.D., Someya, S., Panzer, K., Wohlgemuth, S., Hofer, T., Hacker, T.A., Seo, A.Y., Sullivan, Y., Jobling, W.A., Morrow, J., Van Remmen, H., Sedivy, J.M., Yamasoba, T., Tanokura, M., Saupe, K.W., Weindruch, R., Leeuwenburgh (2005). Mitochondrial DNA mutations, oxidative stress and apoptosis in mammalian aging. Science, 309, 481-484.
View publication via DOI: DOI:10.1126/science.1112125

Gonzalez, A.A., Kumar, R., Mulligan, J.D., Davis, A.J., Weindruch, R., & Saupe, K.W. (2004). Metabolic adaptations to fasting and chronic caloric restriction in heart, muscle, and liver do not include changes in AMPK activity. Am. J. Physiol. Endocrinol. Metab., 287(5), E1032-7.
View publication via DOI: DOI:10.​1152/​ajpendo.​00172.​2004

Bruns, C.M., Baum, S.T., Colman, R.J., Eisner, J.R., Kemnitz, J.W., Weindruch, R., & Abbott, D.H. (2004). Insulin resistance and impaired insulin secretion in prenatally androgenized male rhesus monkeys. J. Clin. Endocrinol. Metab., 89(12), 6218-23.
View publication via DOI: DOI:10.1210/jc.2004-0918

Blanc, S., Schoeller, D., Kemnitz, J., Weindruch, R., Colman, R., Newton, W., Wink, K., Baum, S., & Ramsey, J. (2003). Energy expenditure of rhesus monkeys subjected to 11 years of dietary restriction. J. Clin. Endocrinol. Metab., 88, 16-23.
View publication via DOI: DOI:10.1210/jc.2002-020405

Atwood, C.S., Barzilai, N., Bowen, R.L., Brown-Borg, H.M., Jarrard, D.F., Fu, V.X., Heilbronn, L.K., Ingram, D.K., Ravussin, E., Schwartz, R.S., & Weindruch, R. (2003). Pennington scientific symposium on mechanisms and retardation of aging. Experimental Gerontology, 38(10), 1217-1226.
View publication via DOI: DOI:10.1016/j.exger.2003.08.005

Lee, C.K., Allison, D.B., Brand, J., Weindruch, R., & Prolla, T.A. (2002). Transcriptional profiles associated with aging and middle age-onset caloric restriction in mouse hearts. Proc. Natl. Acad. Sci. USA, 99, 14988-14993.
View publication via DOI: DOI:10.1073/pnas.232308999

Kayo, T., Allison, D.B., Weindruch, R., & Prolla, T.A. (2001). Influences of aging and caloric restriction on the transcriptional profile of skeletal muscle from rhesus monkeys. Proc. Natl. Acad. Sci. USA, 98, 5093-5098.
View publication via DOI: DOI:10.1073/pnas.081061898

Weindruch, R., Keenan, K.P., Fernandes, G., Feuers, R.J., Floyd, R.A., Halter, J.B., Ramsey, J.J., Richardson, A.G., & Spindler, S.R. (2001). Caloric restriction mimetics: Metabolic interventions. J. Gerontol. Biol. Sci., 56A, 20-33.

Lopez, M.E., Van Zeeland, N.L., Dahl, R.D., Weindruch, R., & Aiken, J.M. (2000). Cellular phenotypes of age-associated skeletal muscle mitochondrial abnormalities in rhesus monkeys. Mutation Research, 452, 123-138.
View publication via DOI: DOI:10.1016/S0027-5107(00)00059-2

Ramsey, J.J., Colman, R.J., Binkley, N.C., Christensen, J.D., Gresl, T.A., Kemnitz, J.W., & Weindruch, R. (2000). Dietary restriction and aging in rhesus monkeys: The University of Wisconsin Study. Exp. Gerontol., 35, 1131-1149.

Hall, D.M., Oberley, T.D., Moseley, P.M., Oberley, L.W., Weindruch, R., & Kregel, K.C. (2000). Caloric restriction improves thermotolerance and reduces hyperthermia-induced cellular damage in old rats. FASEB J., 14, 78-86.

Zainal, T.A., Oberley, T.D., Allison, D.B., Szweda, L.I., & Weindruch, R. (2000). Caloric restriction of rhesus monkeys lowers oxidative damage in skeletal muscle. FASEB J., 14, 1825-1836.
View publication via DOI: DOI:10.1096/fj.99-0881com

Lee, C.K., Weindruch, R., & Prolla, T.A. (2000). Gene expression profile of the aging brain in mice. Nat. Genet., 25, 294-297.
View publication via DOI: DOI:10.1038/77046

Ramsey, J.J., Harper, M.E., & Weindruch, R. (2000). Restriction of energy intake, energy expenditure, and aging. Free Radic. Biol. Med., 29, 946-968.
View publication via DOI: DOI:10.1016/S0891-5849(00)00417-2

Pugh, T.D., Oberley, T.D., & Weindruch, R. (1999). Dietary intervention at middle age: caloric restriction but not dehydroepiandrosterone sulfate increases lifespan and lifetime cancer incidence in mice. Cancer Res., 59, 1642-1648.

Lee, C.K., Klopp, R.G., Weindruch, R., & Prolla, T.A. (1999). Gene expression profile of aging and its retardation by caloric restriction. Science, 285, 1390-1393.
View publication via DOI: DOI:10.1126/science.285.5432.1390

Ripple, M.O., Henry, W.F., Schwarze, S.R., Wilding, G., & Weindruch, R. (1999). Effect of antioxidants on androgen-induced AP-1 and NF-kappa B DNA-binding activity in prostate carcinoma cells. J. Natl. Cancer Inst., 91, 1227-1232.

Weindruch, R., & Sohal, R.S. (1997). Caloric intake and aging. New Engl. J. Med., 337, 986-994.

Weindruch, R. (1996). Caloric restriction and aging. Scientific American, 1, 46- 52.

Sohal, R.S., & Weindruch, R. (1996). Oxidative stress, caloric restriction, and aging. Science, 273, 59-63.
View publication via DOI: DOI:10.1126/science.273.5271.59

Weindruch, R., & Walford, R.L. (1988). The retardation of aging and disease by dietary restriction. Springfield IL: C.C. Thomas.

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