Please note: many of the publications offer PDF versions for download. These copies are available to bona fide academics who wish to download single copies of the articles for personal use, and not for commercial gain.

Databases and Index

The following files contain details of my publications in databases for common referencing software packages that can be directly downloaded. These include full publication details and the paper abstracts.

  1. Reference Manager 12 database (.rmx) - Download
  2. Reference Manager 12 database (.rmd) - Download
  3. EndNote database (.ciw) - Download
  4. Text file (compatible with most referencing software) - Download
  5. Index
    The attached file is a word document containing an index of all my publications. You can download this to search for a particular publication or publications on a particular theme etc. To get the associated pdf make a note of the publication's number and then scroll through the following to find the appropriate numbered publication. - Download

Primary Publications - Refereed Journals

  • {517}

    Wanless S., Harris M., SPEAKMAN, J.R., Newell M., Daunt F. (2018 : in press)

    A community-wide decline in the importance of lesser sandeels in seabird chick diet at a North Sea colony

    Marine Ecology Progress series

  • {516}

    Milajerdi, A., Djafarian, K., Bidar, S.S. and SPEAKMAN, J.R. (2018)

    Pre- and post-diagnosis body mass index and heart failure mortality: A dose-response meta-analysis of observational studies reveals greater risk of being underweight than being overweight

    Obesity Reviews

  • {515}

    Kagya-Agyemang, J.K., Vaanholt, L.M., Hambly, C., Król, E., Mitchell, S.E. and SPEAKMAN, J. R. (2018: in press)

    Limits to sustained energy intake XXVIII: Beneficial effects of high dietary fat on lactation performance in mice

    Journal of Experimental Biology

  • {514}

    Hu, S.M., Wang, L., Yang, D.B., Li, L., Togo, J., Wu, Y.G., Liu, Q.S., Li, B.G., Li, M., Wang, G.L., Zhang, X.Y., Li, J.B., Xu, Y.C., Couper, E., Whittington-Davies, A., Mazidi, M., Luo, L.J., Wang, S.G., Douglas, A. and SPEAKMAN, J.R. (2018: in press)

    Dietary fat, but not dietary protein or carbohydrate (sucrose), regulates energy intake and causes adiposity in mice

    CELL metabolism

  • {513}

    SPEAKMAN, J.R., Loos, R.J.F., O’Rahilly, S., Hirschhorn, J.N. and Allison, D.B. (2018: in press)

    GWAS for BMI: a treasure trove of fundamental insights into the genetic basis of obesity.

    International Journal of Obesity

  • {512}

    Nachvak, S.M., Pasdar, Y., Pirsaheb, S., Darbandi, M., Niazi, P., Mostafai, R. and SPEAKMAN, J.R. (2018).

    Effects of Ramadan on food intake, glucose homeostasis, lipid profiles and body composition

    European Journal of Clinical Nutrition

  • {511}

    Mazidid, M. and SPEAKMAN, J.R. (2018)

    The association of fast food and full service restaurant densities with mortality from cardiovascular disease and stroke, and the prevalence of diabetes

    Journal of the American Heart Association

  • {510}

    Green, C.L., Soltow, Q.A., Mitchell, S.E., Derous, D., Wang, Y., Chen, L., Han, J.D.J., Promislow, D.E.L., Lusseau, D., Douglas, A., Jones, D.P. and SPEAKMAN, J.R. (2018 in press).

    The effects of graded levels of calorie restriction: XIII. Global metabolomics screen reveals graded changes in circulating amino acids, vitamins and bile acids in the plasma of C57BL/6 mice.

    Journals of Gerontology series A: Biological Sciences and Medical Sciences

  • {509}

    Mazidi, M. and SPEAKMAN, J.R. (2018)

    The Impact of Obesity and Ozone on the Association between PM2.5 and CVD and Stroke Mortality

    Journal of the American Heart Association

  • {508}

    Fontana, L., Mitchell, S.E., Wang, B.S., Tosti, V., van Vliet, T., Veronese, N., Bertozzi, B., Early, D.S., Maissan, P., SPEAKMAN, J.R. and Demaria, M. (2018)

    The effects of graded caloric restriction XII: Comparison of mouse to human impact on cellular senescence in the colon

    Aging Cell

  • {507}

    Vaanholt, L.M., Duah, O.A., Balducci, S., Mitchell, S.E., Hambly, C. and SPEAKMAN, J.R. (2018)

    Limits to sustained energy intake XXVII: trade-offs between first and second litters in lactating mice support the ecological context hypothesis.

    Journal of Experimental Biology

  • {506}

    Wang, G.L., Cao, M.X., Sauciuvenaite, J., Bissland, R., Hacker, M., Hambly, C., Vaanholt, l.M., Niu, CQ., Faries, M.D. and SPEAKMAN, J.R. (2018)

    Different impacts of resources on opposite sex ratings of physical attractiveness by males and females

    Evolution and Human behaviour 39: 220-225.

  • {505}

    Wilson, G., Lucas, D., Hambly, C., SPEAKMAN, J.R., Morton, J.P. and Close, G.L. (2018)

    Energy expenditure in professional flat jockeys using doubly-labelled water during the racing season: implications for body weight management.

    European Journal of Sport Science 18: 235-242.

  • {504}

    Wang, G.L., Ekeleme-Egedigwe, C.A., El Hamdouchi, A., Sauciuvenaite, J., Bissland, R.M., Djafarian, K., Ojiambo, R., Ramuth, H., Holasek, S., Lackner, S., Diouf, A., Hambly, H., Vaanholt, L.M., Cao, M.X., Hacker, M., Kruger, H.S., Seru, T., Faries, M.D. and SPREAKMAN, J.R. (2018)

    Beauty and the body of the beholder: raters’ body mass index has only limited association with ratings of attractiveness of the opposite sex

    Obesity 26: 522-530

  • {503}

    Lessan, N., Saadane, I., Alkaaf, B., Hambly, C., Buckley, A., Finer, N., SPEAKMAN, J.R., and Barakat, M.T. (2018)

    The effects of Ramadan fasting on activity and energy expenditure

    American Journal of Clinical Nutrition 107:54–61

  • {502}

    Medin, A.C., Carlsen, M.H., Hambly, C., SPEAKMAN, J.R., Stroheimer, S and Andersen, L.F (2017: in press)

    The validity of a web-based food frequency questionnaire assessed by doubly labelled water and multiple 24-hour recalls

    British Journal of Nutrition 118: 1106-07

  • {501}

    SPEAKMAN, J.R. (2017)

    Why lipostatic set-point systems are unlikely to evolve.

    Molecular metabolism

  • {500}

    Moatt, J.P., Hambly, C., Heap, E., Kramer, A., Moon, F., SPEAKMAN, J.R., and Walling, C.A. (2017: in press)

    Body macronutrient composition is predicted by lipid and not protein content of the diet.

    Ecology and Evolution

  • {499}

    Melanson, E., Swibas, T., Khort, W., Catenacci, V., Creasy, S., Plasqui, G., Wouters, L., SPEAKMAN, J.R. and Berman, E. (2017: in press)

    Validation of the doubly labeled water method using off-axis integrated cavity output spectroscopy and isotope ratio mass spectrometry

    American Journal of Physiology – Endocrinology and metabolism.

  • {498}

    Chusyd, D.E., Brown, J.L., Hambly, C., Johnson, M.S., Morfeld, K., Patki, A., SPEAKMAN, J.R., Allison, D.B. and Nagy, T.R. (2017: in press)

    Adiposity and reproductive cycling status in zoo African elephants.


  • {497}

    Zheng, Q.T., Lin, J., Huang, J.J., Zhang, X.Y., Zhang, R., Cao, X.W., Hambly, C., Zhang, H.Y., Ji, Q.T., Wang, X., Qin, G.S., SPEAKMAN, J.R., Wang, Y.F., Jin, W.Z., and Zhao, J.G. (2017: in press)

    Reconstitution of functional UCP1 using CRISPR/Cas9 in the white adipose tissue of pigs decreases fat deposition and improves thermogenic capacity

    Proceedings of the National Academy of the United States

  • {496}

    Qasim, A., Turcotte, M., de Souza, R.J., Samaan, M.C., Champredon, M., Dushoff, J.,

    SPEAKMAN, J.R. and Meyre, D. (2017: in press)

    On the origin of obesity: identifying the biological, environmental, and cultural drivers of genetic risk among human populations

    Obesity Reviews

  • {495}

    SPEAKMAN, J.R. (2017)

    The evolution of body fatness: trading off disease and predation risk

    Journal of Experimental Biology

  • {494}

    Johns, D.W., Marchant, T.A., Fairhurst, G.D., SPEAKMAN, J.R. and Clark, R.G. (2017: in press)

    Biomarker of burden: Feather corticosterone reflects energetic expenditure and allostatic overload in captive waterfowl.

    Functional Ecology

  • {493}

    Riek, A., Brinkmann, L., Gauly, M., Perica, J., Ruf, T., Arnold, W., Hambly, C., SPEAKMAN, J.R. and Gerken, M. (2017)

    Seasonal changes in energy expenditure, body temperature and activity patterns in llamas (Lama glama).

    Scientific reports 7: 7600

  • {492}

    Li, L., Li. B.G., Li, M., Niu, C.Q., Wang, G.L., Li, T., Krol, E., Jin, W.Z. and SPEAKMAN, J.R. (2017)

    Brown adipocytes can display a mammary basal myoepithelial cell phenotype in vivo.

    Molecular metabolism

  • {491}

    Derous, D., Mitchell, S.E., Green, C.L., Wang, Y.C., Han, J.D.J, Chen, L.N., Promislow, D.E.L., Lusseau, D., Douglas, A. and SPEAKMAN, J.R. (2017)

    The Effects of Graded Levels of Calorie Restriction: XI. Evaluation of the main hypotheses underpinning the life extension effects of CR using the hepatic transcriptome

    Aging-US 9:

  • {490}

    Mazidi, M. and SPEAKMAN, J.R. (2017)

    Ambient particulate air pollution (PM2.5) is associated with the ratio of type 2 diabetes to obesity.

    Scientific reports

  • {489}

    Derous, D., Mitchell, S.E., Green, C.L., Wang, Y.C., Han, J.D.J, Chen, L.N., Promislow, D.E.L., Lusseau, D., Douglas, A. and SPEAKMAN, J.R. (2017)

    The Effects of Graded Levels of Calorie Restriction: X. Transcriptomic Responses of Epididymal Adipose Tissue

    Journal of Gerontology: Biological Sciences

  • {488}

    Mazidi, M. and SPEAKMAN, J.R. (2017)

    Higher densities of fast food and full service restaurants are not associated with obesity


    American Journal of Clinical Nutrition 106: 603-613

    This paper was subject of an editorial in the same issue

    Cummins, S., Clary, C. and Sharek, M. (2017) Enduring challenges in estimating the effect of food environment on obesity. Am J Clin Nutr106: 445-446.

    Our response is available online at

    SPEAKMAN, J.R. and Mazidi, M. (2017)

    Food environments and obesity – can’t see the fat for the restaurants?


  • {487}

    Wang, G.L., Li, B.G., Zhang, X.Y., Niu, C.Q., Li, J.B., Li, L., and SPEAKMAN, J.R. (2017).

    No seasonal variation in Physical activity of Han Chinese living in Beijing.

    International Journal of Behavioural Nutrition and Physical Activity 14: 48

  • {486}

    Nilaweera, K.N., Cabrera-Rubio, R., SPEAKMAN, J.R., O’ Connor, P.M., McAuliffe, A., Guinane, C.M., Lawton, E.M., Crispie, F., Aguilera, M., Stanley, M., Boscaini, S., Joyce, S., Melgar, S., Cryan, J.F. and Cotter, P.D. (2017)

    Whey protein-effects on energy balance link the intestinal mechanisms of energy absorption with adiposity and hypothalamic neuropeptide gene expression

    American Journal of Physiology: Endocrinology and metabolism 313: E1-11

  • {485}

    Mohammadi, H., Nidar, S.S., Kazemi, F., Mohseni, A., SPEAKMAN, J.R. and Djafarian, K. (2017: in press).

    Association of single nucleotide polymorphisms in the first intron of the Fat mass and obesity
    associated (FTO) gene with obesity risk in Asians: a meta-analysis

    Progress in Nutrition

  • {484}

    Lin, J., Cao, C.W., Tao, C., Ye, R.C., Dong, M., Zheng, Q.T., Wang, C., Jiang, X.X., Qin, G.S., Yan, C.G., Li, K., SPEAKMAN, J.R., Wang, Y.F., Jin, W.Z., and Zhao, J.G. (2017).

    Cold adaptation in pigs depends on UCP3 in beige adipocytes.

    Journal of Molecular Cell Biology

  • {483}

    Anderson, L., Orme. P., Naughton, R.J., Close, G.L., Milsom, J., Rydings, D., O'Boyle, A., Di Michele, R., Louis, J., Hambly, C., SPEAKMAN, J.R., Morgans, R., Drust, B. and Morton, J.P. (2017)

    Energy intake and expenditure of professional soccer players of the English premier league: evidence of carbohydrate periodization.

    Int. J. Sports Nutrition and Exercise Metabolism. 4: 1-25.

  • {482}

    Yuan, ZQ, Wu, R., Liu, X.M., Sun, G.J., Chen, H., Ma, J., Dong, M., Peng, S.Y., Wang, J.Q., Ding, J.Q., Li, D.H., SPEAKMAN, J.R., Ning, G. and Jin, W.Z. (2017: in press)

    DJ-1 maintains energy and glucose homeostasis by regulating the function of brown adipose tissue.

    Cell Discovery

  • {481}

    Green, C.L., Mitchell, S.E., Derous, D., Wang, Y.C., Han, J.D.J., Chen, L., Promislow, D.E.L., Lusseau, D., Douglas, A. and SPEAKMAN, J.R. (2017)

    The effects of graded levels of calorie restriction: IX. Global metabolomics screen reveals modulation of carnitines, sphingolipids and bile acids in the liver of C57BL/6 mice.

    Aging Cell 16: 529-540

  • {480}

    Mitchell, S.E., Tang, Z.H., Kerbois, C., Delville, C., Derous, D., Green, C.L., Wang, Y.C., Han, J.D.J., Chen, L., Douglas, A., Lusseau, D., Promislow, D.E.L. and SPEAKMAN, J.R. (2017)

    The effects of graded levels of calorie restriction: VIII. impact of short term calorie and protein restriction on basal metabolic rate in the C57BL/6 mouse.

    Oncotarget 8: 17453-17474.

  • {479}

    Mazidi, M., Caravatto, P.P de, SPEAKMAN, J.R. and Cohen, R. (2017)

    Mechanisms of surgery of weight related diseases: the potential role of bile acids.

    Obesity Surgery 27: 826-836.

  • {478}

    Pettett, C.E., Johnson, P.J., Moorhouse, T.P., Hambly, C., SPEAKMAN, J.R. and MacDonald, D.W. (2016: in press)

    Daily energy expenditure in the face of predation: Hedgehog energetics in arable landscapes

    Journal of Experimental Biology

  • {477}

    Da Boit, M., Sibson, R., Sivasubramaniam, S., Meakin, S.J., Greig, C.A., Aspden, R.M., Thies, R., Jeromson, S., Hamilton, D.L., SPEAKMAN, J.R., Hambly, C., Mangoni, A.A., Preston, T., and Gray, S.R. (2016: in press)

    Sex-differences in the effect of fish oil supplementation on the adaptive response to resistance exercise training in older people: a randomized control trial

    American Journal of Clinical Nutrition

  • {476}

    Liu, X.Y., Yang, D.B., Xu, Y.C., Gronning, M.O.L., Zhang, F., Wang, D.H., SPEAKMAN, J.R. (2016)

    Photoperiod induced obesity in the Brandt’s vole (Lasiopodomys brandtii): a model of ‘healthy obesity’?

    Disease models and mechanisms 9: 1357-1366

  • {475}

    Jenniard-Du-Dot, T., Trites, A, Arnould, J.P.Y., SPEAKMAN, J.R. and Guinet, C. (2016: in press)

    Diving, transiting and resting metabolic rates at sea can be estimated from time-activity budgets in free-ranging marine mammals.

    Ecology and Evolution

  • {474}

    Al Jothery, A.H., Vaanhol, L.M., Mody, N., Amous, A., Lykkesfeld, J., Bünger, L., Hill, W.G., Mitchell, S.E. Allison, D.B. and SPEAKMAN, J.R. (2016)

    Oxidative costs of reproduction in mouse strains selected for different levels of food intake and which differ in reproductive performance

    Scientific reports 36353

  • {473}

    Somaye, Y., Mahmoud, K., Alireza, E., SPEAKMAN, J.R., Shemirani, F., Mahdiyeh, N.N., Vida, B. and Djafarian, K. (2016: in press)

    Metabolic Syndrome Patients Have Lower Levels of Adropin When Compared With Healthy Overweight/Obese and Lean Subjects.

    American Journal of Mens Health 2016. pii: 1557988316664074.

  • {472}

    Bagheri, M., SPEAKMAN, J.R., Shemirani, F., and Djafarian, K. (2016: in press)

    Renal Cell Carcinoma risk and obesity: A dose-response meta-analysis reveals another potential paradox within a paradox

    International Journal of Obesity

  • {471}

    Jenniard-Du-Dot, T., Trites, A, Arnould, J.P.Y., SPEAKMAN, J.R. and Guinet, C. (2016: in


    Flipper strokes can predict energy expenditure and locomotion costs in free-ranging northern and Antarctic fur seals

    Scientific reports

  • {470}

    Wang, G.L. and SPEAKMAN, J.R. (2016)

    Analysis of signatures of selection at single nucleotide polymorphisms (SNPs) associated with body mass index (BMI) does not support the ‘Thrifty gene’ hypothesis.

    Cell Metabolism


    The ‘thrifty gene hypothesis’ suggests genetic susceptibility to obesity arises because of positive selection for alleles favored fat deposition and survival during famines. We used public domain data to locate signatures of positive selection based on derived allele frequency, genetic diversity, long haplotypes and differences between populations at single nucleotide polymorphisms (SNPs) identified in genome wide association studies (GWAS) for Body Mass Index (BMI). We used SNPs near the lactase (LCT), SLC24A5 and SLC45A2 genes as positive controls, and 120 randomly-selected SNPs as negative controls. We found evidence for positive selection (p<0.05) at nine out of 115 BMI SNPs. However, five of these involved positive selection for the protective allele (i.e. for leanness). The widespread absence of signatures of positive selection, combined with selection favoring leanness at some alleles, does not support the idea that obesity provided a selective advantage to survive historical periods of famine, or other selective advantage.

  • {469}

    Zhao, Z.J. Yang, D.B., Li, L. Chi, Q.S., Hambly, C. and SPEAKMAN, J.R. (2016 in press)

    Limits to sustained energy intake XXV: milk energy output and thermogenesis in Swiss mice lactating at thermoneutrality.

    Scientific reports

  • {468}

    SPEAKMAN, J.R. and Heidari-Bakavoli, A. (2016: in press)

    Type 2 diabetes, but not obesity, prevalence is positively associated with ambient temperature.

    Scientific reports 6:30409

    Cold exposure stimulates energy expenditure and glucose disposal. If these factors play a significant role in whole body energy balance, and glucose homeostasis, it is predicted that both obesity and type 2 diabetes prevalence would be lower where it is colder. Previous studies have noted connections between ambient temperature and obesity, but the direction of the effect is confused. No previous studies have explored the link of type 2 diabetes to ambient temperature. We used county level data for obesity and diabetes prevalence across the mainland USA and matched this to county level ambient temperature data. Average ambient temperature explained 5.7% of the spatial variation in obesity and 29.6% of the spatial variation in type 2 diabetes prevalence. Correcting the type 2 diabetes data for the effect of obesity reduced the explained variation to 26.8%. Even when correcting for obesity, poverty and race, ambient temperature explained 12.4% of the variation in the prevalence of type 2 diabetes, and this significant effect remained when latitude was entered into the model as a predictor. When obesity prevalence was corrected for poverty and race the significant effect of temperature disappeared. Enhancing energy expenditure by cold exposure will likely not impact obesity significantly, but may be useful to combat type 2 diabetes.

  • {467}

    Jeanniard-du-Dot, T., Trites, A.W., Arnould, J.P.Y., SPEAKMAN, J.R. and Guinet, C. (2016: in press)

    Accelerometers can measure total and activity-specific energy expenditure in free-ranging marine mammals only if linked to time-activity budgets.

    Functional Ecology

  • {466}

    Brinkman, L., Gerken, M., Hambly, C., SPEAKMAN, J.R. and Riek, A. (2016: in press)

    Thyroid hormones correlate with field metabolic rate in ponies (Equus caballus ferus).

    Journal of Experimental Biology

  • {465}

    Vaanholt, L.M., Lane, J.E., Garner, B. and SPEAKMAN, J.R. (2016: in press)

    Partitioning the variance in calorie restriction induced weight and fat loss in outbred mice


  • {464}

    Stothart, M., Elliott, K.H., Wood, T., Hatch, S.A. and SPEAKMAN, J.R. (2016)

    Counting calories in cormorants: dynamic body acceleration predicts daily energy expenditure in pelagic cormorants

    Journal of Experimental Biology

  • {463}

    Morehen, J.C., Bradley, W.J., Clarke, J., Twist, C., Hambly, C., SPEAKMAN, J.R., Morton, J.P. and Close, G. (2016)

    The assessment of total energy expenditure during a 14-day ‘in season’ period of professional rugby league players using the doubly-labelled water method.

    International Journal of Sports Nutrition and Exercise metabolism

  • {462}

    Golzarand, M., Shabobidar, S., Koochakpoor, G. SPEAKMAN, J.R. and Djafarian, K (2016)

    Effect of vitamin D3 supplementation on blood pressure in adults: an updated meta-analysis.

    Nutrition Metabolism and Cardiovascular diseases

  • {461}

    Joly-Amado, A., Serraneau, K., Evsikova, C. SPEAKMAN, J.R., Gordon, M. and Morgan, D. (2016).

    Metabolic changes over the course of aging in a mouse model of tau deposition

    Neurobiology of aging 44:62-73

  • {460}

    Gamo, Y., Bernard, A., Troup, C., Munro, F., Derrer, K., Jeannesson, N., Campbell, A., Gray, H., Miller, J., Dixon, J., Mitchell, S.E., Hambly, C., Vaanholt, L.M., and SPEAKMAN, J.R. (2016)

    Limits to sustained energy intake XXIV: impact of suckling behaviour on the body temperatures of lactating female mice

    Scientific reports 9: e25665

  • {459}

    Derous, D., Mitchell, S.E., Green, C.L., Wang, Y.C., Han, J.D.J., Chen, L., Promislow, D.E.L., Lusseau, D., SPEAKMAN, J.R. and Douglas, A. (2016)

    The Effects of Graded Levels of Calorie Restriction: VII. Topological Rearrangement of Hypothalamic Aging Networks

    Aging-US 8: 917-32

  • {458}

    SPEAKMAN, J.R. and Hambly, C. (2016)

    Doubly-labelled water: some old things to remember and some new things to consider.

    Comparative Biochemistry and Physiology A

  • {457}

    SPEAKMAN, J.R., Mazidi, M. and Mitchell, S.E. (2016)

    Calories or Protein? The effect of dietary restriction on lifespan in rodents is due to calories alone.

    Experimental Gerontology

  • {456}

    Jafarnejad, S., Shab-Bidar, S., SPEAKMAN, J.R., Parastui, K., Daneshi-Maskooni, M. and Djafarian, K.(2016: in press)

    Probiotics reduce the risk of antibiotic associated diarrhea in adults (18 – 64 y) but not the elderly (>65 y): a meta-analysis.

    Nutrition in Clinical practice

  • {455}

    Mitchell, S.E., Derous, D., Green, C.L., Chen, L., Han, J.D.J., Wang, Y.C., Promislow, D.E.L., Lusseau, D., Douglas, A. and SPEAKMAN, J.R. (2016)

    The Effects of Graded Levels of Calorie Restriction: V. Impact of Short-term Graded Calorie Restriction on physical activity in the C57BL/6 mouse.


  • {454}

    Krol, E., Douglas, A., Tocher, D.R., Crampton, V.O., SPEAKMAN, J.R., Secombes, C.J. and Martin, S.A.M. (2016).

    Differential responses of the gut transcriptome to plant protein diets in farmed salmon.

    BMC Genomics 17: 156

  • {453}

    Derous, D., Mitchell, S.E., Green, C.L., Chen, L., Han, J.D.J., Wang, Y.C., Promislow, D.E.L., Lusseau, D., SPEAKMAN, J.R. and Douglas, A. (2016)

    The Effects of Graded Levels of Calorie Restriction: VI. Impact of Short-term Graded Calorie Restriction on Transcriptomic Responses of the Hypothalamic Hunger and Circadian Signaling Pathways


  • {452}

    Burke, L.K., Doslikova, B., D’Agostino, G., Greenwald-Yarnell, M., Georgescu, T., Chianese, R., Martinez de Morentin, P.B., Ogunnowo-Bada, E., Cansell, C., Torres, L.V., Garfield, A.S., Schoute, J.S., Lam, D.D., SPEAKMAN, J.R., Rubinstein, M., Low, M.J., Rochford, J.J., Myers, M.G., Evans, M.L., and Heisler, L.K. (2016: in press)

    Sex difference in Physical activity, energy expenditure and obesity driven by a sub-population of hypothalamic POMC neurons

    Molecular Metabolism 5: 245-252.

  • {451}

    Sadowska, E.T., Krol, E., Chrzascik, K.M., Rudolf, A., SPEAKMAN, J.R., and Koteja, P. (2016)

    Limits to sustained energy intake XXIII. Does heat dissipation capacity limit energy budgets of lactating bank voles?

    Journal of Experimental Biology 219: 805-815

  • {450}

    Lipina, C., Vaanholt, L.M., Davidova, A., Storey-Gordon, E., Mitchell, S.E., Hambly, C., Irving, A.J.,SPEAKMAN, J.R. and Hundal, H.S. (2016)

    CB1 receptor blockade counters age-induced insulin resistance and Metabolic Dysfunction.

    Aging Cell

  • {449}

    Angel, L.P., Wells, M.R., Malagon, M.R., Tew, E., SPEAKMAN,, J.R. and Arnould, J.P.Y (2015)

    Sexual size dimorphism and body condition in the Australasian Gannet

    Plos ONE 10: e0142653

  • {448}

    Vaanholt, L.M., Milne, A., Zheng, Y., Hambly, C., Mitchell, S.E., Valencak, T.G., Allison, D.B. and SPEAKMAN, J.R. (2015: in press)

    Oxidative Costs of Reproduction: Oxidative Stress in Mice Fed Standard and Low Antioxidant Diets

    Physiology and Behaviour 154: 1-7

  • {447}

    SPEAKMAN, J.R., Blount, J.D., Bronikowski, A.M., Buffenstein, R., Isaksson, C., Kirkwood, T.B.L., Monaghan, P., Ozanne, S.E., Beaulieu, M., Briga, M., Carr, S.K., Christensen, L.L., Cochemé, H.M., Cram, D.L., Dantzer, B., Harper, J.M., Jurk, D., King, A., Noguera, J.C., Salin, K., Sild, E., Simons, M.J.P., Smith, S., Stier, A., Tobler, M., Vitikainen, E., Peaker, M. and Selman, C. (2015: in press)

    Oxidative stress and life histories: unresolved issues and current needs

    Ecology and Evolution


    Life history theory concerns the trade-offs that mould the patterns of investment by animals between reproduction, growth and survival. It is widely recognised that physiology plays a role in the mediation of life history trade-offs, but the details remain obscure. As life history theory concerns aspects of investment in the soma that influence survival, understanding the physiological basis of life histories is related, but not identical, to understanding the process of ageing. One idea from the field of ageing that has gained considerable traction in the area of life histories is that life history trade-offs may be mediated by free-radical production and oxidative stress. We outline here developments in this field and summarise a number of important unresolved issues that may guide future research efforts. The issues are as follows. First, different tissues and macromolecular targets of oxidative stress respond differently during reproduction. The functional significance of these changes, however, remains uncertain. There is a need for studies that link oxidative stress measurements to functional outcomes, such as survival. Second, measurements of oxidative stress are often highly invasive or terminal. Terminal studies of oxidative stress in wild animals, where detailed life history information is available, cannot generally be performed without compromising the aims of the studies that generated the life history data. There is a need therefore for novel non-invasive measurements of multi-tissue oxidative stress. Third, laboratory studies provide unrivalled opportunities for experimental manipulation but may fail to expose the physiology underpinning life history effects, because of the benign laboratory environment. Fourth, the idea that oxidative stress might underlie life history trade-offs does not make specific enough predictions that are amenable to testing. Moreover, there is a paucity of good alternative theoretical models on which contrasting predictions might be based. Fifth, there is an enormous diversity of life history variation to test the idea that oxidative stress may be a key mediator. So far we have only scratched the surface. Broadening the scope may reveal new strategies linked to the processes of oxidative damage and repair. Finally, understanding the trade-offs in life histories and understanding the process of ageing are related but not identical questions. Scientists inhabiting these two spheres of activity seldom collide, yet they have much to learn from each other.

  • {446}

    Wang, G.L., Djafarian, K., Egedigwe, C.A., Asmaa, E.H., Ojiambo, R., Ramuth, H., Wallner-Liebermann, S.J., Lackner, S., Diouf, A., Sauciuvenaite, J., Hambly, C., Vaanholt, L.M., Faries, M.D. and SPEAKMAN, J.R. (2015)

    The relationship of body fatness to female attractiveness.

    PeerJ 3: e1155


    Aspects of the female body may be attractive because they signal evolutionary fitness. Greater body fatness might reflect greater potential to survive famines, but individuals carrying larger fat stores may have poor health and lower fertility innon-famine conditions. A mathematical statistical model using epidemiological data linking fatness to fitness traits, predicted a peaked relationship between fatness and attractiveness (maximum at body mass index (BMI) = 23.2 to 24.8 depending on ethnicity and assumptions). Participants fromthree Caucasian populations (Austria,Lithuania and the UK), three Asian populations (China, Iran and Mauritius) and four African populations (Kenya,Morocco, Nigeria and Senegal) rated attractiveness of a series of female images varying in fatness (BMI) and waist to hip ratio (WHR).
    There was an inverse linear relationship between physical attractiveness and body fatness or BMI in all populations. Lower body fat was more attractive, down to at least BMI = 19. There was no peak in the relationship over the range we studied in
    any population.WHRwas a significant independent but less important factor, which was more important (greater r2) in African populations. Predictions based on the fitness model were not supported. Raters appeared to use body fat percentage (BF%)
    and BMI as markers of age. The covariance of BF% and BMI with age indicates that the role of body fatness alone, as a marker of attractiveness, has been overestimated.

    This paper recieved worldwide press coverage when realesed in August 2015

  • {445}

    Lusseau, D., Mitchell, S.E., Barros, C., Derous, D., Green, C., Chen, L., Han, J.D.J., Wang, Y.C., Promislow, D.E.L., Douglas, A. and SPEAKMAN, J.R. (2015)

    The effects of graded levels of caloric restriction, IV: Non-linear change in behavioural phenotype of male C57BL/6 mice in response to short-term graded caloric restriction.

    Scientific reports 5: 13198


    Animals have to adjust their activities when faced with caloric restriction (CR) to deal with reduced energy intake. If CR is pronounced, allostasis can push individuals into alternate physiological states which can result in important health benefits across a wide range of taxa. Here we developed a new
    approach to determine the changes in behavioural phenotype associated with different levels of CR. We exposed C57BL/6 male mice to graded CR (from 0 to 40%) for three months and defined their behavioural phenotype using hidden Markov models of their movement and body temperature. All 40% CR mice exhibited a state-shift in behavioural phenotype and only some exposed to 30% CR did. We show for the first time that mice changed their activity characteristics rather than changed their activities. This new phenotyping approach provides an avenue to determine the mechanisms linking CR to healthspan.

  • {444}

    Monarca, R.I., Mathias, M. Da Luz and SPEAKMAN, J.R. (2015)

    Behavioural and physiological responses of wood mice (Apodemus sylvaticus) to experimental manipulations of predation and starvation risk

    Physiology and Behaviour 149: 331-339


    Body weight and the levels of stored body fat have fitness consequences. Greater levels of fat may provide protection
    against catastrophic failures in the food supply, but theymay also increase the risk of predation. Animals may therefore regulate their fatness according to their perceived risks of predation and starvation: the starvation– predation trade-off model. We tested the predictions of this model in wood mice (Apodemus sylvaticus)
    by experimentally manipulating predation risk and starvation risk.We predicted that under increased predation risk individuals would lose weight and under increased starvation risk they would gain it. We simulated increased predation risk by playing the calls made by predatory birds (owls: Tyto alba and Bubo bubo) to the
    mice. Control groups included exposure to calls of a non-predatory bird (blackbird: Turdus merula) or silence. Mice exposed to owl calls at night lost weight relative to the silence group, mediated via reduced food intake, but exposure to owl calls in the day had no significant effect. Exposure to blackbird calls at night also resulted in weight loss, but blackbird calls in the day had no effect. Mice seemed to have a generalised response to bird calls at night irrespective of their actual source. This could be because in the wild any bird calling at night will be a predation risk, and any bird calling in the day would not be, because at that time the mice would normally be resting, and hence not exposed to avian predators. Consequently, mice have not evolved to distinguish different types of call but only to respond to the time of day that they occur. Mice exposed to stochastic 24 h starvation
    events altered their behaviour (reduced activity) during the refeeding days that followed the deprivation periods
    to regain the lostmass. However, they only marginally elevated their food intake and consequently had reduced body weight/fat storage compared to that of the control unstarved group. This response may have been constrained by physiological factors (alimentary tract absorption capacity) or behavioural factors (perceived risk of predation). Overall the responses of the mice appeared to provide limited support for the starvation–
    predation trade-off model, and suggest that wood mice are much more sensitive to predation risk than they are to starvation risk.

  • {443}

    Hambly, C. and SPEAKMAN, J.R. (2015)

    Mice that gorged during dietary restriction increased foraging related behaviors and differed in their macronutrient preference when released from restriction.

    PeerJ 3: e1091


    Caloric restriction (CR) can trigger gorging behavior.We examined macronutrient choice and behavior in mice that gorged during restriction compared to restricted non-gorgers and controls. Fifty MF1 male mice were restricted to 75% of ad-libitum food intake (FI), while ten controls were fed ad-lib. Body mass (BM) and FI were measured two and 24-h after food inclusion over 14-days. ‘Gorging’ mice were defined as those which ate over 25% of their daily FI in 2-h. The top 11 gorgers and the lowest 9 gorgers, along with 10 controls, had their behavior analysed during restriction, and were then provided with an unrestricted food choice, consisting of three diets that were high in fat, protein or carbohydrate. During restriction gorgers ate on average 51% of their daily FI in the 2-h following food introduction while the non-gorgers ate only 16%. Gorgers lost significantly more BM than non-gorgers possibly due to an increased physical activity linked to anticipation of daily food provision. Controls and non-gorgers spent most of their time sleeping. After restriction, both gorgers and non-gorgers were hyperphagic until their lost weight was regained. All 3 groups favoured high fat food. Gorgers and non-gorgers had a significantly greater high carbohydrate diet intake than controls, and gorgers also had a significantly greater high protein diet intake than non-gorgers and controls. On unrestricted food, they did not continue to gorge, although they still had a significantly greater 2-h FI than the other groups. Elevated protein intake may play an important role in the recovery of the lost lean tissue of gorgers after restriction.

  • {442}

    Vaanholt, L.M., Mitchell, S.E., Sinclair, R.E. and SPEAKMAN, J.R. (2015)

    Increased food anticipatory activity and differential expression of melanocortin3 receptor (MC3R) and Dopamine receptor 2 (D2) in mice that are resistant versus prone to diet-induced weight loss

    Hormones and behaviour 73: 83-93


    Diet-induced weight loss varies considerably between individuals, but the mechanisms driving these individual differences remain largely unknown. Herewe investigatedwhether key neuropeptides involved in the regulation of energy balance or reward systemswere differentially expressed in mice that were prone or resistant to caloric restriction (CR) induced weight loss. Mice (n=30males and n=34 females) were fed 70% of their ownbaseline
    ad libitum intake for 25 days, after which their brains were collected and expression of various neuropeptides were investigated and compared between the 10male and 10 female mice that showed the greatest (highweight loss,HWL) or lowestweight loss (LWL) (n=40 in total).HWLmice showed a differential neuropeptide profile to LWL in both sexes, characterised by increased expression of neuropeptide Y (NPY), agouti-related peptide (AgRP), leptin receptor (ObRb), and melanocortin 3 receptor (MC3R) in the arcuate nucleus. No changes in the expression of fat mass and obesity related gene (FTO) or suppressor of cytokine signalling 3 (Socs3) were observed. Levels of dopamine D2 receptor were decreased in the nucleus accumbens in HWL compared to LWL mice. HWL mice showed a stronger increase in food anticipatory activity (FAA) in response to CR than
    LWL mice. These results indicate that the mice prone to diet-induced weight loss experienced greater hunger, potentially driving their elevated FAA.

  • {441}

    Monarca, R.I., Mathias, M. da Luz, Wang, D.H. and SPEAKMAN, J.R. (2015)

    Predation risk modulates diet induced obesity in male C57BL/6 mice.



    Objective: In this study we aimed to examine the behavioural and physiological changes induced by experimentally varying the risk of predation, in male mice fed on a high fat diet. In particular, we aimed to assess if the risk of being predated modulates the body weight gain, providing an ecological context for the obesity resistance observed in many species of small mammals.

    Methods: Body weight, food intake, physical activity and core body temperature of 35 male C57BL/6 mice were monitored for twenty days, whilst feeding on high fat diet. A third of the animals were exposed to elevated risk of predation through exposure to the sounds of nocturnal predatory birds and these were compared to animals exposed to a neutral noise or silence.

    Results: Male mice exposed to predation risk had significantly lower weight gain than the neutral or silent groups. Reduced of food intake and increased physical activity were the main proximal factors explaining this effect. The risk of predation also induced changes in boldness.

    Conclusions: Our study provides evidence supporting the role of predation risk on body weight gain of small mammals, sustaining the influence of environmental factors on body weight regulation mechanisms.

  • {440}

    Nie, Y.G.*, SPEAKMAN, J.R.*, Wu, Q.*, Zhang, C.L., Hu, Y.B., Xia, M.H., Li, Y., Hambly, C., Wang, L., Wei, W., Zhang, J.G. and Wei, F.W. (2015)

    The bamboo eating giant panda has low daily energy expenditure

    Science 349: 171-175

    *Joint first authors


    The carnivoran giant panda has a specialized bamboo diet, to which its alimentary tract is poorly adapted. Measurements of daily energy expenditure across five captive and three wild pandas averaged 5.2 megajoules (MJ)/day, only 37.7%of the predicted value (13.8 MJ/day). For the wild pandas, themean was 6.2 MJ/day, or 45%of themammalian expectation. Pandas achieve this exceptionally low expenditure in part by reduced sizes of several vital organs and low physical activity. In addition, circulating levels of thyroid hormones thyroxine (T4) and triiodothyronine (T3) averaged 46.9 and 64%, respectively, of the levels expected for a eutherian mammal of comparable size. A giant panda–niquemutation in the DUOX2 gene, critical for thyroid hormone synthesis, might explain these low thyroid hormone levels. A combination of morphological, behavioral, physiological, and genetic adaptations, leading to low energy expenditure, likely enables giant pandas to survive on a bamboo diet.

    See also

    'Inside Science' review of the paper in the same issue

    Vignieri, S. (2015) Laid back bamboo eater. Science 349: This week in Science

    and Armitage, H (2015) Science shot. AAAS Video report on the paper. Why are pandas so lazy?

    SPEAKMAN, J.R. (2015) Why pandas are so chilled. The clue is in the bamboo. The Conversation. 10 July 2015. This article was published in The Conversation to coincide with the papers release. It was viewed 53,200 times in the first week after release.

    SPEAKMAN, J.R. (2015) Pandas are cool – its official. Newton

  • {439}

    Polyviou, T.P., Pitsiladis, Y.P., Celis-Morales, C.A., Brown, B., SPEAKMAN, J.R. and Malkova, D. (2015)

    The Effects of Hyperhydrating Supplements Containing Creatine and Glucose on Plasma Lipids and Insulin Sensitivity in Endurance Trained Athletes

    Journal of Amino Acids

  • {438}

    McManus, B., Korpela, R., SPEAKMAN,J.R., O’Connor, P., Cryan, J.F., Cotter, P.D. and Nilaweera, K.N. (2015)

    Bovine serum albumin as the dominant source of protein reduces subcutaneous fat mass, plasma leptin, and plasma corticosterone in high fat fed C57/BL6 mice

    British Journal of Nutrition 114: 654-662.

  • {437}

    Mitchell, S.E., Delville, C., Konstantopedos, P., Hurst, J., Derous, D., Green, C., Chen, L.N., Han, J.D.J., Wang, Y.C., Promislow, D.E.L., Lusseau, D., Douglas, A., and SPEAKMAN, J.R. (2015).

    The effects of graded levels of calorie restriction: III. Impact of short term calorie and protein restriction on mean daily body temperature and torpor use in the C57BL/6 mouse.

    Oncotarget 6: 18314-18337


    A commonly observed response in mammals to calorie restriction (CR) is reduced body temperature (Tb). We explored how the Tb of male C57BL/6 mice responded to graded CR (10 to 40%), compared to the response to equivalent levels of protein
    restriction (PR) over 3 months. Under CR there was a dynamic change in daily Tb over the first 30–35 days, which stabilized thereafter until day 70 after which a further decline was noted. The time to reach stability was dependent on restriction level.
    Body mass negatively correlated with Tb under ad libitum feeding and positively correlated under CR. The average Tb over the last 20 days was significantly related to the levels of body fat, structural tissue, leptin and insulin-like growth factor-1. Some
    mice, particularly those under higher levels of CR, showed periods of daily torpor later in the restriction period. None of the changes in Tb under CR were recapitulated by equivalent levels of PR. We conclude that changes in Tb under CR are a response only to the shortfall in calorie intake. The linear relationship between average Tb and the level of restriction supports the idea that Tb changes are an integral aspect of the lifespan effect.

  • {436}

    Mitchell, S.E., Delville, C., Konstantopedos, P., Hurst, J., Derous, D., Green, C., Chen, L., Han, J.D.J., Wang, Y.C., Promislow, D.E.L., Lusseau, D., Douglas, A., and SPEAKMAN, J.R. (2015).

    The effects of graded levels of calorie restriction: II. Impact of short term calorie and protein restriction on circulating hormone levels, glucose homeostasis and oxidative stress in male C57BL/6 mice.

    Oncotarget 6: 23213-23237


    imiting food intake attenuates many of the deleterious effects of aging, impacting upon healthspan and leading to an increased lifespan. Whether it is the overall restriction of calories (calorie restriction: CR) or the incidental reduction in macronutrients such as protein (protein restriction: PR) that mediate these effects is
    unclear. The impact of 3 month CR or PR, (10 to 40%), on C57BL/6 mice was compared to controls fed ad libitum. Reductions in circulating leptin, tumor necrosis factor-α and insulin-like growth factor-1 (IGF-1) were relative to the level of CR and individually
    associated with morphological changes but remained unchanged following PR. Glucose tolerance and insulin sensitivity were improved following CR but not affected by PR. There was no indication that CR had an effect on oxidative damage, however CR
    lowered antioxidant activity. No biomarkers of oxidative stress were altered by PR. CR significantly reduced levels of major urinary proteins suggesting lowered investment in reproduction. Results here support the idea that reduced adipokine levels, improved insulin/IGF-1 signaling and reduced reproductive investment play important roles in the beneficial effects of CR while, in the short-term, attenuation of oxidative damage
    is not applicable. None of the positive effects were replicated with PR.

  • {435}

    Araujo-Soares, V., Sniehotta, F.F., Laing, C.M., Gellert, P., Jackson, D.M. & SPEAKMAN, J.R. (2015)

    Social cognitions measured in 4 to 6 year olds are predictive of objectively measured physical activity

    Psychology and Health


    Objective: This study tested whether social cognitions from the Reasoned Action Approach (RAA) can be reliably measured in a sample of primary school children (aged 4 to 6 years)
    and whether these RAA measures are predictive of physical activity.

    Design: Longitudinal Observational study with repeated measures over 6 months. Measures: RAA variables were
    measured using a novel choice paradigm between physically active and sedentary pastimes. Relative Reinforcing value, covariates (sex, age, deprivation, BMI), and physical activity by
    accelerometry (Primary outcome) were also measured.

    Results: RAA cognitions could be measured with acceptable retest-reliability and discriminant validity and social norms regarding parents and teachers (but not friends), were correlated with physical activity and change in physical activity when relative reinforcing value and other covariates were controlled for.

    Conclusions: RAA cognitions can be reliably measured in primary school children aged 4-6 and RRA measures appear useful in understanding children’s activity
    choices and may potentially inform future interventions.

  • {434}

    Rizzolo, D.J., Schmutz, J.A. and SPEAKMAN, J.R. (2015)

    Fast and efficient: post-natal growth and energy expenditure in an Arctic breeding waterbird, the red-throated loon (Gavia stellata).

    The Auk 132: 657-670


    Environmental conditions can exert a strong influence on the growth and energy demands of chicks. We hypothesized that postnatal growth in a cold, aquatic environment would require a high level of energy metabolism in semiprecocial Red-throated Loon (Gavia stellata) chicks. We measured body-mass growth and daily energy expenditure (DEE) of freeranging chicks in the Arctic. We used daily gains in body mass and DEE to estimate daily metabolizable energy (DME, kJ day1) and total metabolizable energy (TME, kJ chick1). Chicks gained body mass quickly, with a logistic growth rate constant 57% greater than the allometric prediction, yet were at only 60% of adult body mass at fledging. Males grew at a rate similar to that of females but for a slightly longer duration and so reached an asymptotic body mass 23%
    greater, and tarsus length 8% longer, than that of females. Chick growth performance was similar between first- and second-hatched chicks within broods of 2, which suggests that food availability was not limited. DEE increased in proportion to body mass, and DME peaked at 1,214 kJ day1 on day 25 posthatching. Over the average 49-day postnatal period, TME was 49.0 MJ, which is within the range of error of the allometric prediction. Parents provided
    58.6 MJ as food to meet this energy requirement. Given this chick energy requirement and the range of energy content of prey observed in the chick diet, selecting prey with higher energy content would greatly reduce adult provisioning effort. Red-throated Loon chicks did not have a high postnatal energy requirement, but rather grew quickly and fledged at a small size—with the effect of reducing the length of the postnatal period and, consequently, parental energy investment in chicks.

  • {433}

    Vaanholt, L.M., Sinclair, R., Mitchell, S.E and SPEAKMAN, J.R. (2015)

    Factors influencing individual variability in high fat diet induced weight gain in out-bred MF1 mice.

    Physiology and Behaviour 144:46-155


    Easy access to high-energy palatable foods has been suggested to have contributed to the world-wide obesity epidemic. However, within these ‘obesogenic’ environments many people manage to remain lean. Mice also show variability in their weight gain responses to high-fat diet (HFD) feeding and their weight loss responses to calorically restricted (CR) feeding. In this study we investigated which factors contribute to determining susceptibility
    to HFD-induced obesity in mice, and whether the responses in weight gain on HFD are correlated with the responses to CR. One-hundred twenty four mice were exposed to 30% CR for 28 days followed by a 14 day recovery period, and subsequent exposure to 60% HFD for 28 days. Responses in various metabolic factors were
    measured before and after each exposure (body mass; BM, body composition, food intake; FI, resting metabolic rate; RMR, physical activity, body temperature and glucose tolerance; GT). Weight changes on HFD ranged from−1 to 26%, equivalent to−0.2 g to 10.5 g in absolute mass.Multiple regression models showed that fat free mass (FFM) of the mice before exposure to HFD predicted 12% of the variability inweight gain on HFD (p b 0.001). Also, FI during the firstweek of HFD feeding predicted 20% of the variability in BM and fat mass (FM) gain 4 weeks later. These data may point to a role for the reward system in driving individual
    differences in FI andweight gain.Weight gain on theHFD was significantly negatively correlated to weight loss on CR, indicating that animals that are poor at defending against weight gain on HFD, were also poor at defending against CR-induced weight loss. Changes in FM and FFM in response to HFD or CRwere not correlated however.

  • {432}

    Derakshanian, H., Shab-Bidar, S., SPEAKMAN, J.R., Nadimi, H. and Djafarian, K. (2015)

    Vitamin D and diabetic nephropathy: a systematic review and meta-analysis.

    Nutrition 31: 1189-1194


    Objective: There has been a long history documenting the use of different vitamin D derivatives as therapy for renal diseases. However, to our knowledge, there is no comprehensive assessment of the relation between vitamin D deficiency and risk for diabetic nephropathy (DN). Additionally, the effect of vitamin D supplementation on DN is still unclear. The aim of this meta-analysis was to assess these issues by pooling together the results from cross-sectional studies and clinical trials.

    Methods: A systematic literature search of PubMed, Scopus, and Google Scholar was conducted, ending in September 2014. For cross-sectional studies, odds ratio was used as a measure of the
    association between vitamin D status and risk for DN; for clinical trials, mean and SD of the main outcome (urine albumin-to-creatinine ratio [UACR]) in intervention and placebo groups were
    considered for analysis.

    Results: The final selected articles were published between 2009 and 2014. In all, 3700 and 219
    patients were enrolled in observational and interventional studies, respectively. The pooled odds ratio from six cross-sectional studies was 1.80 (95% confidence interval [CI], 1.25–2.59; P ¼ 0.002),
    indicating a significant inverse association between serum vitamin D status and risk for nephropathy in patients with diabetes. However, the pooled data of UACR levels in clinical trials
    suggested no significant change following vitamin D supplementation (17.98; 95% CI, –35.35 to
    71.32; P ¼ 0.51).

    Conclusion: This meta-analysis showed the higher risk for nephropathy in vitamin D–deficient patients with diabetes. Pooling the results of available clinical trials after vitamin D supplementation did not support causality in this association.

  • {431}

    Djafarian, K., SPEAKMAN, J.R., Stewart, J. and Jackson, D.M.
    Body composition and basal metabolic rate in pre-school children:
    no sex difference.

    Journal of Nutritional Sciences and Dietetics.

  • {430}

    Liu, X.M., Wang, S.P., You, Y.L., Meng, M.H., Zheng, Z.J., Dong, M., Lin, J., Zhao, Q.W., Zhang, C.H., Yuan, X.X., Hu, T., Liu, L.Q., Huang, Y.Y., Zhang, L., Wang, D.H., Zhan,J.C., SPEAKMAN, J.R. and Jin, W.Z. (2015)

    Brown adipose tissue transplantation reverses obesity in the Ob/Ob mouse.



    Increasing evidence indicates that brown adipose tissue (BAT) transplantation enhances whole body energy metabolism in a mouse model of diet-induced obesity. However, it remains unclear
    whether BAT also has such beneficial effects on genetically obese mice. To address this issue, we transplanted BAT (trBAT) from C57/BL6 mice into the dorsal subcutaneous (SUB) region of age and sex matched leptin deficient Ob/Ob mice. Interestingly, BAT transplantation led to a significant reduction of body weight gain with increased oxygen consumption and decreased total body fat
    mass, resulting in improvement of insulin resistance and liver steatosis. In addition, BAT transplantation increased the level of circulating adiponectin, while it reduced the levels of circulating
    free triiodothyronine (T3) and thyroxine (T4) which regulate thyroid hormone sensitivity in peripheral tissues. BAT transplantation also increased3 adrenergic receptor and fatty acid oxidation related gene expression in SUB and epididymal (EP) white adipose tissue. Accordingly, BAT transplantation
    increased whole-body thermogenesis. Taken together our results demonstrate that BAT transplantation may reduce obesity and its related diseases by activating endogenous BAT.

  • {429}

    Merkling, T., Welcker, J., Hewison, M., Hatch, S.A., Kitaysky, A., SPEAKMAN, J.R. , Danchin, E., and Blanchard, P. (2015)

    Identifying the selective pressures underlying offspring sex-ratio adjustments

    Behavioral Ecology26: 916-925


    Sex allocation theory predicts that parents should bias offspring sex according to the costs and benefits associated with producing
    either sex in a given context. Accurately interpreting sex-ratio biases, therefore, requires a precise identification of these
    selective pressures. However, such information is generally lacking. This may partly explain the inconsistency in reported sex
    allocation patterns, especially in vertebrates. We present data from a long-term feeding experiment in black-legged kittiwakes
    (Rissa tridactyla) that allowed us to increase investment capacity for some breeding pairs. Previous findings showed that these
    pairs then overproduced sons compared with control parents. Here, our aim was to test the underlying assumptions of the 2
    appropriate sex allocation models for our context: the “cost of reproduction hypothesis” and the “Trivers–Willard hypothesis.”
    The former assumes a sex difference in rearing costs, whereas the latter assumes a difference in fitness returns. 1) Independent
    of feeding treatment, rearing sons was energetically more demanding for parents (as revealed by higher energy expenditure
    and higher baseline corticosterone levels) than rearing daughters, thereby corroborating the underlying assumption of the “cost
    of reproduction hypothesis.” 2) Evidence supporting the assumptions of the “Trivers–Willard hypothesis” was less convincing. Overall, our results suggest that drivers of parental sex allocation decisions are probably more related to offspring sex-specific energetic costs than to their future reproductive success in our study species. Assessing the adaptive value of sex-ratio biases
    requires precise investigation of the assumptions underlying theoretical models, particularly as long as the mechanisms involved in sex-ratio manipulation remain largely unknown.

  • {428}

    Mitchell, S.E., Tang, Z.H., Kerbois, C., Delville, C., Konstantopedos, P., Bruel, A. Derous, D., Green, C., Aspden, R.M., Goodyear, S.R., Chen, L.N., Han, J.J.D., Wang, Y.C., Promislow, D.E.L., Lusseau, D., Douglas, A. and SPEAKMAN, J.R. (2015)

    The effects of graded levels of calorie restriction: I. Impact of short term calorie and protein restriction on body composition in the C57BL/6 mouse.

    Oncotarget 6: 15902-15930


    Faced with reduced levels of food, animals must adjust to the consequences of the shortfall in energy. We explored how C57BL/6 mice withdrew energy from different body tissues during three months of food restriction at graded levels up to 40% (calorie restriction: CR). We compared this to the response to equivalent
    levels of protein restriction (PR) without a shortfall in calories. Under CR there was a dynamic change in body mass over 30 days and thereafter it stabilized. The time to reach stability was independent of the level of restriction. At the end of three
    months whole body dissections revealed differential utilization of the different tissues. Adipose tissue depots were the most significantly utilized tissue, and provided 55.8 to 60.9% of the total released energy. In comparison, reductions in the sizes of
    structural tissues contributed between 29.8 and 38.7% of the energy. The balance was made up by relatively small changes in the vital organs. The components of the alimentary tract grew slightly under restriction, particularly the stomach, and this was
    associated with a parallel increase in assimilation efficiency of the food (averaging 1.73 %). None of the changes under CR were recapitulated by equivalent levels of PR.

  • {427}

    Bagheri, M., SPEAKMAN, J.R., Shabbidar, S., Kazemi, F., and Djafarian, K. (2015)

    A dose-response meta-analysis of the impact of body mass index on stroke and all cause mortality in stroke patients: a paradox within a paradox.

    Obesity Reviews 16: 416-423


    The obesity paradox is often attributed to fat acting as a buffer to protect individuals in fragile metabolic states. If this was the case, one would predict that the reverse epidemiology would be apparent across all causes of mortality including that of the particular disease state. We performed a dose-response meta-analysis to assess the impact of body mass index (BMI) on all-cause and
    stroke-specific mortality among stroke patients. Data from relevant studies were identified by systematically searching PubMed, OVID and Scopus databases and were analysed using a random-effects dose-response model. Eight cohort studies
    on all-cause mortality (with 20,807 deaths of 95,651 stroke patients) and nine studies of mortality exclusively because of stroke (with 8,087 deaths of 28,627 patients) were evaluated in the meta-analysis. Non-linear associations of BMI with all-cause mortality (P < 0.0001) and mortality by stroke (P = 0.05) were
    observed. Among overweight and obese stroke patients, the risk of all-cause mortality increased, while the risk of mortality by stroke declined, with an increase in BMI. Increasing BMI had opposite effects on all-cause mortality and stroke-specific mortality in stroke patients. Further investigations are needed to examine how mortality by stroke is influenced by a more accurate indicator of
    obesity than BMI.

  • {426}

    Careau, V., Montiglio, P.O., Garant, D., Pelletier, F., SPEAKMAN, J.R., Humphries, M.M. and Reale, D. (2015)

    Energy expenditure and personality in wild chipmunks

    Behavioral Ecology and Sociobiology 69: 653-661


    According to the Bpace-of-life syndrome^ concept,
    slow-fast life-history strategies favored under different ecological
    conditions should lead to co-adaptations between metabolic
    rate and personality traits such as activity, exploration,
    and boldness. Although the relationships between resting metabolic
    rate (RMR) and personality traits have been recently
    tested several times, we still do not know whether personality
    is related to the daily energy expenditure (DEE) of free-living
    individuals in their natural habitat. The objectives of this study
    were to assess the links between RMR, DEE, and two personality
    traits (exploration in an open-field and docility during
    handling) in wild eastern chipmunks (Tamias striatus).
    Using a multivariate mixed model, we found that exploration
    and docility were significantly correlated at the amongindividual
    level, confirming the presence of a behavioral syndrome
    within our population. We also found that exploration,
    but not docility, was negatively correlated with DEE. Hence,
    fast explorers show lower DEE levels than slow explorers,
    independently of RMR and docility. This result adds to an
    increasingly large (and complex) literature reporting the impacts
    of personality traits on the biology, ecology,

  • {425}

    Richardson, A., Fischer, K.E., SPEAKMAN, J.R., de Cabo, R., Mitchell, S.J., Peterson, C.A., Rabinovitch, P., Chiao, Y.A., Taffet, G., Miller, R.A., Rentería, R.C., Bower, J., Ingram, D.K., Ladiges, W.C., Ikeno, Y., Sierra, F. and Austad, S.N. (2015: in press)

    Measures of Healthspan As Indices of Aging in Mice – a Recommendation

    Journals of Gerontology B

  • {424}

    Berman, E.S.F., Melanson, E.L., Swibas, T., Snaith, S.P., and SPEAKMAN, J.R. (2015)

    Inter- and intra-individual correlations of background abundances of 2H, 18O and 17O in human urine and implications for DLW measurements

    European Journal of Clinical Nutrition 69: 1091-1098

  • {423}

    Saboori, S., Shab-Bidar, S., SPEAKMAN, J.R. Rad, E.Y., and Djafarian, K. (2015)

    Effect of vitamin E supplementation on serum C-reactive protein level: a meta-analysis of randomized controlled trials

    European Journal of Clinical Nutrition8: 867-873

  • {422}

    SPEAKMAN, J.R. (2015)

    The ‘fat mass and obesity related’ (FTO) gene: mechanisms of impact on obesity and energy balance

    Current Obesity Reports 4(1). DOI: 10.1007/s13679-015-0143-1

  • {421}

    Verant, M.L., Meteyer, C.U., SPEAKMAN, J.R., Cryan, P.M., Lorch, JM., and Blehert, D.S. (2014)

    White-nose syndrome initiates a cascade of physiologic disturbances in the hibernating bat host

    BMC Physiology 14:10

  • {420}

    Santoro, S., Green, A.J., SPEAKMAN, J.R. and Figuerola, J. (2015).

    Effects of population density on condition and sex-ratio of glossy ibis.


  • {419}

    McAllen, L., SPEAKMAN, J.R., Cryan, J.F. and Nilaweera, K.N. (2015)

    Whey protein isolate decreases murine stomach weight and intestinal length, and alters expression of Wnt signaling associated genes.

    British Journal of Nutrition 113: 372-379.

  • {418}

    Duarte, L.C. and SPEAKMAN, J.R. (2014)

    Resting metabolic rate is negatively related to lifespan because of a confounding effect of body fatness.

    AGE 36: 9731

  • {417}

    Brinkmann, L., Gerken, M., Hambly, C., SPEAKMAN, J.R. and Riek, A. (2014)

    Saving energy during hard times: Energetic adaptations of Shetland pony mares

    Journal of Experimental Biology 217: 4120-4127

  • {416}

    Szafrańska, P.A., Zub, K., Wieczorek, M., Książek, A. SPEAKMAN, J.R. and Konarzewski, M. (2014)

    Shaving increases daily energy expenditure in free living root voles.

    Journal of Experimental biology 217: 3964-3967

  • {415}

    Dhurandhar, N.V., Schoeller, D.A., Brown, A.W., Heymsfield, S.B., Thomas, D., Sørensen, T.I.A., SPEAKMAN, J.R. Jeansonne, M., Allison, D.B. and the Energy Balance Measurement Working Group (2015)

    Energy Balance Measurement: When Something is Not Better than Nothing

    International Journal of Obesity 39: 1109-1113

    See also follow-up letter

    Dhurandhar, N.V., Schoeller, D.A., Brown, A.W., Heymsfield, S.B., Thomas, D., Sørensen, T.I.A., SPEAKMAN, J.R. Jeansonne, M., Allison, D.B (2015)

    Authors response to LTE for Energy Balance Measurement: When Something is Not Better than Nothing.

    International Journal of Obesity 39: 1175-1176

  • {414}
    • Al Jothery, A.H., Krol, E., Hawkins, J., Chetoui, A., Saint-Lambert, A., Gamo, Y., Shaw, S.C., Valencak, T., Bunger, L., Hill, W.G., Vaanholt, L.M., Hambly, C., and SPEAKMAN, J.R. (2014)

    Limits to sustained energy intake XXII: Reproductive performance of two selected mouse lines with different thermal conductance.

    Journal of Experimental Biology

    Brief Summary: lines long term selected for high and low food intake differ in their thermal conductance. We tested if this leads to elevated reproductive performance in line with the predictions of the heat dissipation limits theory. It does.


    Maximal sustained energy intake (SusEI) appears limited, but the factors imposing the limit are disputed. We studied reproductive performance in two lines of mice selected for high and low food intake (MH and ML, respectively), and known to have large differences in thermal conductance (29% higher in the MH line at 21°C). When these mice raised their natural litters, their metabolisable energy intake significantly increased over the first 13 days of lactation and then reached a plateau. At peak lactation, MH mice assimilated on average 45.3 % more energy than ML mice (222.9±7.1 and 153.4±12.5 kJ day−1, N=49 and 24, respectively). Moreover, MH mice exported on average 62.3 kJ day−1more energy as milk than ML mice (118.9±5.3 and 56.6±5.4 kJ day−1, N=subset of 32 and 21, respectively). The elevated milk production of MH mice enabled them to wean litters (65.2±2.1 g) that were on average 50.2% heavier than litters produced by ML mothers (43.4±3.0 g), and pups that were on average 27.2% heavier (9.9±0.2 and 7.8±0.2 g, respectively). Lactating mice in both lines had significantly longer and heavier guts compared to non-reproductive mice. However, inconsistent with the central limit hypothesis, the ML mice had significantly longer and heavier intestines than MH mice. An experiment where the mice raised litters of the opposing line demonstrated that lactation performance was not limited by offspring growth capacity. Our findings are consistent with the idea that the SusEI at peak lactation is constrained by the capacity of the mothers to dissipate body heat.

  • {413}

    Scantlebury, D.M., Mills, M.G.L., Wilson, R.P., Wilson, J.W., Mills, M.E.J., Durant, S.M., Bennett, N.C., Bradford, P., Marks, N.J. and SPEAKMAN, J.R. (2014)

    Flexible energetics of cheetah hunting strategies provide resistance against kleptoparasitism

    Science 346: 79-81


    Population viability is driven by individual survival, which in turn depends on individuals balancing energy budgets. As carnivores may function close to maximum sustained power outputs, decreased food availability or increased activity may render some populations energetically vulnerable. Prey theft may compromise energetic budgets of mesopredators, such as cheetahs and wild dogs, which are susceptible to competition from larger carnivores. We show that daily energy expenditure (DEE) of cheetahs was similar to size-based predictions and positively related to distance traveled. Theft at 25% only requires cheetahs to hunt for an extra 1.1 hour per day, increasing DEE by just 12%. Therefore, not all mesopredators are energetically constrained by direct competition.
    Other factors that increase DEE, such as those that increase travel, may be more important for population viability.

    See also inside Science review of this paper in the same issue

    Laundre, J.W. (2014) How large predators manage their energy.Science 346: 33

  • {412}

    Elliott, K.H., Chivers, L., Bessey, L., Gaston, A.J., Hatch, S.A., Kato, A., Osborne, O., Ropert-Coudert, Y., SPEAKMAN, J.R. and Hare, J.F. (2014)

    Against the wind: windscapes shape seabird instantaneous energy costs but adult behaviour buffers impact on offspring.

    Movement Ecology 2:17

  • {411}

    Piper, M.D.W., Selman, C., SPEAKMAN, J.R. and Partridge, L. (2014)

    Using doubly-labeled water to measure energy expenditure in an important small ectotherm

    Drosophila melanogaster

    Journal of Genetics and Genomics

  • {410}

    Welcker, J., SPEAKMAN, J.R., Elliott, K.H., Hatch, S.A., and A. Kitaysky (2014)

    Resting and daily energy expenditures during reproduction are adjusted in opposite directions in free-living birds

    Functional ecology



    1. Reproduction is energetically expensive, and daily energy expenditure (DEE) often peaks during the period of rearing young. The “potentiation” hypothesis predicts that high DEE needs to be sustained by a corresponding up-regulation of metabolic machinery, thus a concomitant increase of the resting metabolic rate (RMR) is expected. Alternatively, the “compensation” hypothesis predicts that DEE and RMR are regulated independently and animals may maintain low RMR to maximize the energy available for reproduction. This might particularly be the case if DEE was limited, e.g. by extrinsic food supply or intrinsic physiological factors.
    2. We tested these hypotheses in free-living seabirds by manipulating their energy demands (experimentally reduced or increased brood size) and supplies (providing supplemental food), and simultaneously measuring their DEE and RMR (by the doubly-labeled water method and an indirect hormonal proxy, respectively).
    3. In support of the ‘compensation’ hypothesis, metabolic rates were adjusted independently and in opposite directions with an increase of DEE and a decrease of the hormonal proxy for RMR in individuals rearing young compared to birds with removed broods. DEE of unfed birds appeared to be limited and supplemental feeding did not allow DEE to exceed this limitation.
    4. We propose that a reduction of the resting metabolism is a strategy to increase allocation of energy to reproduction when DEE is constrained, and that this constraint is unlikely to be related to food supply.
  • {409}

    Bagheri, M., Ansari, S., Sotoudeh, G., Mahmoudi, M., SPEAKMAN, J.R. and Djafarian, K. (2015)

    Serum ghrelin levels and gender-related indices of body composition in prepubertal children: a cross-sectional study.

    European Journal of Clinical Nutrition 68:

  • {408}

    Nachvak, S.M., Neyestani, T.R., Mahboob, S.A., Sabour, S., Keshawarz, S.A. and SPEAKMAN, J.R. (2014)

    α-tocopherol supplementation reduces biomarkers of oxidative stress in children with Down syndrome: a randomized controlled trial.

    European Journal of Clinical Nutrition

    Brief summary: Oxidative stress has been implicated in the cognitive decline in Down syndrome. Consistent with this Down syndrome children had greater oxidative damage to DNA than their siblings. Supplementation with vitamin E reduced their oxidative stress levels.


    Background: Down syndrome (DS) is the most common human chromosomal abnormality. It is characterized by mental retardation and several metabolic disturbances, including elevated oxidative stress, which may be causally linked. Treatment with dietary antioxidants has been suggested as a potential method to alleviate the oxidative damage and retardation of DS patients, but prior supplementation work has been equivocal.

    Aim: To evaluate the effects of supplementation with antioxidants α-tocopherol and α-lipoic acid (ALA) on oxidative stress biomarkers in DS children.

    Methods: Ninety-three DS children aged 7-15 years from both sexes were randomly allocated to three groups: α-tocopherol (400 IU/d), ALA (100 mg/d) and placebo. The intervention period was four months. A healthy control group consisted 26 non-DS siblings. Serum thiobarbituric acid reactive substances (TBARS) and urinary 8-hydroxydeoxyguanosine (8OHdG) were used as biomarkers of oxidative stress.

    Results: DS children had greater levels of baseline oxidative stress than their siblings. Moreover, males had greater levels of 8OHdG than females (p < 0.001) but there was no significant association between age and biomarkers of oxidative stress. Serum levels of TBARS did not change significantly over time, or relative to placebo. Although urinary 8OHdG concentrations decreased significantly in both α-tocopherol and ALA, groups compared to the baseline levels (p < 0.001), mean final levels of urinary 8OHdG concentrations differed significantly only between α-tocopherol and placebo groups (p< 0.01).

    Conclusion: α-tocopherol supplementation of the diets of DS children may attenuate oxidative stress at the DNA level.

  • {407}

    Müller,M.J., Baracos, V., Bosy-Westphal, A., Dulloo, A., Eckel, J., Fearon, K.C.H., Hall, K.D., Pietrobelli, A., Sørensen, T.I.A., SPEAKMAN, J.R., Trayhurn, P., Visser, M., and Heymsfield, S.B. (2014)

    Functional Body Composition and Related Aspects in Research on Obesity and Cachexia

    Obesity reviews 15: 640-656.

  • {406}

    Qian, P., Li, M., Shi, Y.L., Liu, H.W., SPEAKMAN, J.R. and Wang, D.H. (2014)

    Lipidomics reveals mitochondrial membrane remodelling associated with acute thermoregulation in a rodent with a wide thermoneutral zone.

    Lipids 40: 715-730


    Living in xeric environments, Mongolian gerbils (Meriones unguiculatus) have high physiological flexibility in response to acute temperature change, and have the widest thermoneutral zone (TNZ, 26.5oC-38.9oC) reported among small mammals. At the upper critical temperature (Tuc), body temperatures of gerbils were significantly elevated while metabolic rates were maintained at basal level. In contrast, below the lower critical temperature (Tlc) metabolism was elevated and body temperature stable. We predicted that mitochondrial membrane remodeling may play an important role during acute thermoregulation of gerbils, as it is rapid and energetically inexpensive. Here, acute changes in mitochondrial membrane lipids and related energetic parameters were examined between Tuc (38oC) and Tlc (27oC), and between Tlcand 16oC. Detailed changes in mitochondrial membrane lipids were detected by lipidomic analysis. Compared to 27oC, a decrease in n-6 and zwitterionic phospholipids in muscle mitochondrial membranes was found at 38oC, together with elevated glucose utilization. At 16oC, the fluidity of mitochondrial membrane in both muscle and brown adipose tissue was significantly increased , indicated by a significant increase in the unsaturation index and reduced fatty acid chain lengths of membrane phospholipids. Our results suggest that mitochondrial membrane lipid remodeling may stabilize membrane function when body temperature is elevated, and improve heat production under acute cold exposure. These data therefore emphasize the important role of membrane remodeling during short-term thermoregulation in a non-hibernating endotherm.

  • {405}

    Fletcher, Q.E., SPEAKMAN, J.R., Boutin, S., Lane, J.E., McAdam, A.G., Gorrell, J.C., Coltman, D.W. and Humphries, M.M. (2014)

    Daily energy expenditure during lactation is strongly selected for in a free-living mammal

    Functional Ecology 29: 195-208

  • {404}

    Djafarian, K., SPEAKMAN, J.R., Stewart, J. and Jackson, D.M. (2014)

    Comparison of activity levels measured by a wrist worn accelerometer and direct observation of young children

    Open Journal of Pediatrics 3: 422-427

  • {403}

    SPEAKMAN, J.R. (2014)

    If body fatness is under physiological regulation, then how come we have an obesity epidemic?

    Physiology 29: 88-98.

    This paper was part of a themed issue of 6 papers on

    obesity. The theme was illustrated on the cover of the issue

  • {402}

    Zhang, Z., Hao, C.J., Li, C.G., Zhao, J., Li, X.N., Wei, A.H., Wei, Z.B., He, X., Zhen, X.C., Gao, X., SPEAKMAN, J.R. and Li, W. (2014)

    Mutation of SLC35D3 Causes Metabolic Syndrome by Impairing Dopamine Signaling in Striatal D1 Neurons

    PLOS Genetics 10: e1004124

    Brief Summary Mutation in SLC35D3 gene in mice leads to failure to traffic dopamine receptor 1 from ER to the cell surface of striatum neurons. This leads to diminished physical activity and daily energy expenditure leading to symptoms similar to metabolic syndrome. One in 200 Han Chinese with metabolic syndrome have mutations in SLC34D3 gene.

    Media summary and impact file : download word doc


    Obesity is one of the largest health problems facing the world today. Although twin and family studies suggest about two-thirds of obesity is caused by genetic factors, only a small fraction of this variance has been unraveled. There are still large numbers of genes to be identified that cause variations in body fatness and the associated diseases encompassed in the metabolic syndrome (MetS). A locus near a sequence tagged site (STS) marker D6S1009 has been linked to obesity or body mass index (BMI). However, its genetic entity is unknown. D6S1009 is located in the intergenic region between SLC35D3 and NHEG1. Here we report that the rosmutant mice, harboring a recessive mutation in the Slc35d3 gene, show obesity and MetS and reduced membrane dopamine receptor D1 (D1R) with impaired dopamine signaling in striatal neurons. SLC35D3 is localized to both endoplasmic reticulum (ER) and early endosomes and interacts with D1R. In ros striatal D1 neurons, lack of SLC35D3 causes the accumulation of D1R on the ER to impair its ER exit. The MetS phenotype is reversible by the administration of D1R agonist to the ros mutant. In addition, we identified two mutations in the SLC35D3 gene in patients with MetS, which alter the subcellular localization of SLC35D3. Our results suggest that theSLC35D3 gene,close to the D6S1009 locus, is a candidate gene for MetS, which is involved in metabolic control in the central nervous system by regulating dopamine signaling.

  • {401}

    Valencak, T.G., Wright, P., Weir, A., Mitchell, S.E., Vaanholt, L.M., Hambly, C., Krol, E. and SPEAKMAN, J. R. (2013)

    Limits to sustained energy intake XXI: Effect of exposing the mother, but not her pups, to a cold environment during lactation in mice.

    Journal of Experimental Biology 216: 4326-4333

    doi: 10.1242/jeb.092023

    Brief summary: Lactating female mice able to visit a cold compartment that their offspring could not access had elevated food intake and milk production, but their pups did not grow any faster. The data support the heat dissipation limits model and highlight the problems of inferring female milk production from pup growth rates.

    Chinese summary: 哺乳期雌性小鼠在进入冷暴露的小隔间(他们的后代无法进入)会增加摄食和奶产量,但是他们后代的生长不会加快。这些数据支持散热限制假说模型凸显了女性产奶量与后代生长率的问题。


    The capacity of females to dissipate heat may constrain sustained energy intake during lactation. However, some previous experiments supporting this concept have confounded the impact of temperature on the mothers with the impact on the pups. We aimed to separate these effects in lactating laboratory mice (MF1 strain) by giving the mothers access to cages at 2 ambient temperatures (10 and 21° C) joined by a tube. Food was available only in the cold cage, but females could also choose go to this cage to cool down while pups were housed in the warmer cage. Control animals had access to the same configuration of cages but with both maintained at 21° C. We hypothesised that if females were limited by heat dissipation, alleviating the heat load by providing a cool environment would allow them to dissipate more heat, intake more food, generate more milk and hence wean heavier litters. We measured maternal energy budgets and monitored time courses of core body temperature and physical activity. To minimize the variance in energy budgets all litters were adjusted to 12 (+/- 1) pups. Females in the experimental group had higher energy intake (F1,14= 15.8; p=0.0014), higher assimilated energy (F1,13= 10.7; p=0.006) and provided their pups with more milk (F1,13= 6.65; p=0.03), consistent with the heat dissipation limit theory. Yet, despite keeping demand constant, mean pup growth rates were similar (F 1,13=0.06; p=0.8) thus our data emphasise the difficulties of inferring milk production indirectly from pup growth

    pdf available on JEB web site

    and also here

  • {400}

    SPEAKMAN, J.R. and Garratt, M. (2014)

    Oxidative stress as a cost of reproduction: beyond the simplistic trade-off model.

    Bioessays 36: 93-106.

  • {399}

    Djafarian, K., SPEAKMAN, J.R., Stewart, J. and Jackson, D.M. (2013)

    Familial resemblance of body composition, physical activity and resting metabolic rate in pre-school children.

    Reports of biochemistry and molecular biology 2: 1-15

  • {398}

    Fletcher, Q.E., Landry-Cuerrier, M., Boutin, S., McAdam, A.G., SPEAKMAN, J.R. and Humphries, M.M. (2013)

    Reproductive timing and reliance on hoarded capital resources by lactating red squirrels.

    Oecologia.173: 1203-1215

    doi: 10.1007/s00442-013-2699-3


    Successful reproduction in a seasonal environment can be accomplished with resources that are stored before use (“capital resources”) or resources that are used immediately (“income resources”). Research examining capital versus income resource usage during reproduction has primarily focused on assigning species to positions along a capital-income gradient. Here, we examine the causes and reproductive consequences of among and within-year variation in hoarded capital versus income resource usage by female North American red squirrels (Tamiasciurus hudsonicus) during mid-lactation in a highly seasonal environment. Among-years, the proportion of feeding events that were on capital resources (PROPCAP) averaged 39% during the yearly median mid-lactation periods, but ranged widely between 2% and 100%. In years with earlier parturition dates, females primarily used hoarded capital resources during mid-lactation, whereas in years with later parturition dates, females primarily used income resources during mid-lactation. Within-years, PROPCAP during mid-lactation tended to be greater in early-breeding females than in late-breeding females. Rates of water flux in females during mid-lactation provided further evidence that late-breeding females used more water-rich income resources. The proportion of litters that were partially or completely lost, and the litter mass that lactating females supported, was not influenced by the large among-year differences in hoarded capital resource usage. Red squirrels appear to delay reproduction following years with low cone production to time peak reproductive demands to be late enough to be supported by income resources that only become available later in the season. In conclusion, our results offer a rare example of the capacity of a food-hoarding mammal to support reproduction exploiting a wide range of capital and income resources.

    pdf available at springer website

    and here

  • {397}

    Guo, X.G., SPEAKMAN J.R., Dong, W.G., Men, X.Y., Qian, T.J., Wu, D., Qin, F. and Song WY. (2013)

    Ectoparasitic insects and mites on Yunnan red-backed voles (Eothenomys miletus) from a localized area in southwest China.

    Parasitology Research. 112: 3543-3549


    Ectoparasitic insects and mites on Yunnan redbacked
    voles (Eothenomys miletus) in Dali prefecture,
    Yunnan Province, southwest China, were studied between
    2003 and 2004. In total, 34,389 individuals of 86 species of
    ectoparasitic insects (seven species of fleas and five species
    of sucking lice) and mites (23 species of gamasid mites and 51 species of chigger mites) were collected from 916 individual

    hosts. The diversity of ectoparasites on this single rodent
    species in such a small area was much higher than in previous
    reports, which concerned more host species and greater geographical
    areas. The majority of the ectoparasites were chigger
    mites, which accounted for 59.3 % of the parasite species and
    87.4 % of the individual parasites. Most voles harbored parasites
    with an overall prevalence (P) of 82.5 % and mean
    abundance (MA) of 37.5 parasites per host. The dispersion
    coefficient (C) and patchiness index (m*/m) were used to
    study the spatial patterns of the seven dominant parasite
    species, and all seven had aggregated distributions. The species
    abundance distribution of the ectoparasites on the vole
    was fitted by Preston’s lognormal distribution (R2=0.82), and
    the total expected parasite species was estimated fromthis plot
    as 167 species. Yunnan red-backed voles harbor many ectoparasites as revealed by examination of a large host population. Future field investigations should sample large numbers
    of host individuals to assess ectoparasite populations.

  • {396}

    Elliott, K.H., Vaillant, M.L., Kato, A., Gaston, A.J., Ropert-Coudert, Y., Hare, J.F., SPEAKMAN, J.R. and Croll, D. (2014)

    Age-related variation in energy expenditure of a long-lived bird within the envelope of an energy ceiling.

    Journal of Animal Ecology 83: 136-146.

    doi: 10.1111/1365-2656.12126.


    1. Energy expenditure in wild animals can be limited (i) intrinsically by physiological processes that constrain an animal’s capacity to use energy, (ii) extrinsically by energy availability in the environment and/or (iii) strategically based on trade-offs between elevated metabolism and survival. Although these factors apply to all individuals within a population, some individuals expend more or less energy than other individuals.
    2. To examine the role of an energy ceiling in a species with a high and individually repeatable metabolic rate, we compared energy expenditure of thick-billed murres (Uria lomvia) with and without 17 g handicaps affixed to their backs during a period of peak energy demand (chick-rearing, N = 16). We also compared energy expenditure of unencumbered birds (N = 260) across 8 years exhibiting contrasting environmental conditions and correlated energy expenditure with fitness (reproductive success and survival).
    3. Murres appeared to experience an energy ceiling mediated through behavioural adjustments. Handicapped birds decreased time spent flying/diving and chick provisioning rates such that overall daily energy expenditure remained unchanged across the two treatments. The energy ceiling likely did not reflect energy availability or trade-offs with fitness, as energy expenditure was similar across contrasting foraging conditions and was not associated with reduced survival or increased reproductive success. Across individuals within a species, older murres, with lower future reproductive potential, did invest more in offspring following handicapping. Thus, we found partial support for the trade-off hypothesis, as individuals were able to overcome an energy ceiling over the scale of days, apparently at the cost of current energy reserves at least among older birds, where prospects for future reproduction would be relatively limited.
    4. A meta-analysis comparing responses of breeding animals to handicapping suggests that our results are typical: animals either reduced investment in themselves or in their offspring to remain below an energy ceiling. Across species, whether a handicapped individual invested in its own energy stores or its offspring’s growth was not explained by life-history (future vs. current reproductive potential). Many breeding animals apparently experience an intrinsic energy ceiling.

    pdf available from wiley website

    and here

  • {395}

    Xu,Y.C., Yang, D.B., SPEAKMAN, J.R. and Wang, D.H. (2014)

    Oxidative stress in response to natural and experimentally elevated reproductive effort is tissue dependent

    Functional Ecology 28: 402-410

    doi: 10.1111/1365-2435.12168

    Brief summary: The oxidative stress theory suggests life history trade-offs between reproduction and survival may be due to the oxidative stress caused by reproduction. This paper shows that depending on what tissues you examine the data support or refute the hypothesis.

    Chinese summary: 氧化应激理论表明在繁殖和生存的生活史的权衡可能是因为氧化应激是由于繁殖所导致的,这篇论文表明基于不同组织的研究可能会支持或反驳这一假说。


    1. Oxidative stress is a potential proximal physiological cost of reproduction. Detecting this cost may be performed in several different ways – manipulating reproductive status, correlating natural variations in effort to oxidative stress, or manipulating reproductive effort. Here, we manipulated reproductive status and studied oxidative stress due to natural and experimental variation in reproductive effort in Brandt’s voles (Lasiopodomys brandtii), using a variety of markers and tissues.
    2. We measured markers of oxidative stress in lactating (raising 6 to 8 offspring) and non-reproductive voles (Experiment I) and found a marker of oxidative protection (serum total-superoxide dismutase (SOD) activity) was reduced, and a marker of oxidative damage (protein carbonyls) was increased, in the serum, in lactating compared with non-reproductive voles. However, protein carbonyls in the liver were lower in lactating compared with non-reproductive voles, consistent with increased liver SOD activity. Lipid damage (malondialdehyde (MDA)) in both serum and liver was unrelated to reproductive status.
    3. We compared these markers of oxidative stress between natural large (n ≥ 9) and small litter sizes (n ≤ 5) (Experiment II), and between manipulated large (11-13) and small litter sizes (2-3) (Experiment III) and found that liver MDA and SOD activity was higher in voles with natural large compared to natural small litter sizes, but there were no differences in other markers. There was no effect of litter size on all measures when it was experimentally manipulated.
    4. The effects of reproductive status on oxidative stress were critically dependent on the exact markers and tissues used. The effects of natural variation in reproductive effort suggested there might be an oxidative stress cost associated with large litter sizes, but this effect was not replicated in the experimentally manipulated litters.

    lay summary available here

    supplementary data available here

    see also discussion of issues relating to this paper on research gate

    Click here to go to RG discussion

    pdf available on wiley website

    and here

  • {394}

    Yang, D.B., Xu, Y.C., Wang, D.H. and SPEAKMAN, J.R. (2013)

    Effects of reproduction on immuno-suppression and oxidative damage, and hence support or otherwise for their roles as mechanisms underpinning life history trade-offs, are tissue and assay dependent

    Journal of Experimental Biology 216: 4242-4250

    doi: 10.1242/jeb.092049

    Brief summary: The inferred effects of reproduction on immunosuppression and oxidative damage, and hence support or otherwise for particular physiological mechanisms that underpin life history trade-offs, are critically dependent on the exact markers and tissues used.

    Chinese summary:繁殖对免疫抑制或氧化损伤的的推论性的影响因此支持或者会巩固生活史权衡的特殊的生理机制主要是基于研究所用的标记和组织。


    Life history parameters appear to be traded off against each other, but the physiological mechanisms involved remain unclear. One hypothesis is that potentially energetically costly processes such as immune function, and protection from oxidative stress, may be compromised during reproductive attempts because of selective resource allocation. Lower temperatures also impose energy costs, and hence allocation decisions might be more pronounced when animals are forced to reproduce in the cold. Here, we experimentally tested whether reproduction at different ambient temperatures was associated with elevated oxidative stress and suppressed immune function in Mongolian gerbils (Meriones unguiculatus). Using a variety of different markers for both immune function and oxidative stress we found that some measures of immune function (serum bactericidal capacity and size of the thymus) were significantly suppressed, while some measures of oxidative protection (serum superoxide dismutase (SOD) activity and glutathione peroxidase (GPx) activity) were also reduced, and a marker of oxidative damage (protein carbonyls in serum) was increased, in lactating compared with non-reproductive gerbils. These changes were in line with the selective resource allocation predictions. However, the PHA response and serum total immunoglobulin (IgG) were not suppressed, and other markers of oxidative damage (malondialdehyde MDA (TBARS) and protein carbonyls in the liver) were actually lower in lactating compared with non-reproductive gerbils, consistent with increased levels of SOD activity and total antioxidant capacity (T-AOC) in the liver. These latter changes were opposite the expectations based on resource allocation. Furthermore other measures of protection (GPx levels in the liver and protein thiols in both serum and liver) and damage (MDA (TBARS) in serum) were unrelated to reproductive status. Ambient temperature differences did not impact on these patterns. Collectively, our results indicated that the inferred effects of reproduction on immunosuppression and oxidative damage, and hence support or otherwise for particular physiological mechanisms that underpin life history trade-offs , are critically dependent on the exact markers and tissues used. This may be because during reproduction individuals selectively allocate protection to some key tissues, but sacrifice protection of others.

    see also discussion of issues relating to this paper on research gate

    click here to go to RG discussion

    pdf available from JEB website

    and here

  • {393}

    Gamo, Y., Troup, C., Mitchell, S.E., Hambly, C., Vaanholt, L.M., andSPEAKMAN, J.R. (2013)

    Limits to sustained energy intake XX: body temperatures and physical activity of female mice during lactation

    Journal of Experimental Biology 216: 3751-3761

    doi: 10.1242/jeb.090308


    Lactating animals consume greater amounts of food than non-reproductive animals but energy intake appears to be limited in late lactation. The heat dissipation limit theory suggests that the food intake of lactating mice is limited by the capacity of the mother to dissipate heat. Lactating mice should therefore have high body temperatures (Tb), and changes in energy intake, during lactation, should be reflected in variation in Tb. To investigate these predictions, 26 mice (Mus musculus) were monitored daily throughout lactation for food intake, body mass, litter size and litter mass. After weaning, 21 days postpartum, maternal food intake and body mass were monitored for another 10 days. Maternal activity and Tb were recorded every minute for 23 hours a day using implanted transmitters (vital view). Energy intake increased to a plateau in late lactation (days 13-17). Daily gain in pup mass declined during this same period, suggesting a limit on maternal energy intake. Litter size, and litter mass, were positively related to maternal energy intake and body mass. Activity levels were constantly low, and mice with the largest increase in energy intake at peak lactation had the lowest activity. Tb rose sharply after parturition and the circadian rhythm became compressed within a small range. Tb during the light period increased considerably (1.1oC higher than in baseline), and lactating mice faced chronic hyperthermia, despite their activity levels in lactation being approximately halved. Average Tb increased in relation to energy intake as lactation progressed,, but there was no relationship between litter size or litter mass and the mean Tb at peak lactation. These data are consistent with the heat dissipation limit theory which suggests performance in late lactation is constrained by the ability to dissipate body heat.

    pdf available at JEB website

    as well as here

  • {392}

    Wilson, J.W., Mills, M.G.L., Wilson, R.P., Peters, G., Mills, M.E.J., SPEAKMAN, J.R., Durant, S.M., Bennett, N,C., Marks, N.J. and Scantlebury, M. (2013)

    Cheetahs Acinonyx jubatus balance turn capacity with pace when chasing prey

    Biology Letters 9:20130620

    doi: 10.1098/rsbl.2013.0620


    Predators–prey interactions are fundamental in the evolution and structure of ecological communities. Our understanding, however, of the strategies used in pursuit and evasion remains limited. Here we report on the hunting dynamics of the world’s fastest land animal, the cheetah Acinonyx jubatus. Using miniaturized data loggers, we recorded fine-scale movement, speed and acceleration of free-ranging cheetahs to measure how hunting dynamics relate to different prey chases. Cheetahs attained hunting speeds of up to 18.94 m/s and accelerated up to 7.5 m/s with greatest angular velocities achieved during the terminal phase of the hunt. The interplay between forward and lateral acceleration during chases showed that the total forces involved in speed changes and turning were approximately constant over time but varied with prey type. Thus, rather than a simple maximum speed chase, cheetahs first accelerate to decrease the distance to their prey before reducing speed 5-8 s from the end of the hunt, so as to facilitate rapid turns to match prey escape tactics, varying precise strategy according to prey species. Predator and prey thus pit a fine balance of speed against manoeuvring capability in an evolutionary arms race for survival.

    pdf available from royal society publishing website here

    as well as here

  • {391}

    Selman, C., McLaren, J.S., Collins, A.R., Duthie, G.G. andSPEAKMAN, J.R. (2013)

    Deleterious consequences of antioxidant supplementation on lifespan in a wild-derived mammal.

    Biology Letters 9: 20130432

    doi: 10.1098/rsbl.2013.0432

    Data archived at data dryad.com

    Summary of media coverage and impact word doc

    Brief Summary: Voles fed high doses of vitamin E or Vitamin C had dramatically reduced lifespans compared with unsupplemented individuals, despite reduced lipid peroxidation levels in their livers, suggesting the dietary supplements were reducing oxidative stress.

    Chinese summary: 被饲喂高剂量的维生素E和维生素C的田鼠和那些被饲喂的田鼠相比,其寿命会显著地缩短,尽管降低了它们肝脏中的脂质的过氧化水平,表明膳食的添加物会减少氧化应激。


    While oxidative damage owing to reactive oxygen species (ROS) often increases with advancing age and is associated with many age-related diseases, its causative role in ageing is controversial. In particular, studies that have attempted to modulate ROS-induced damage, either upwards or downwards, using antioxidant or genetic approaches, generally do not show a predictable effect on lifespan. Here, we investigated whether dietary supplementation with either vitamin E (a-tocopherol) or vitamin C (ascorbic acid) affected oxidative damage and lifespan in short-tailed field voles,
    Microtus agrestis. We predicted that antioxidant supplementation would reduce ROS-induced oxidative damage and increase lifespan relative to unsupplemented controls. Antioxidant supplementation for nine months reduced hepatic lipid peroxidation, but DNA oxidative damage to hepatocytes and lymphocytes was unaffected. Surprisingly, antioxidant supplementation significantly
    shortened lifespan in voles maintained under both cold (7+28C)
    and warm (22+28C) conditions. These data further question the predictions of free-radical theory of ageing and critically, given our previous research in mice, indicate that similar levels of antioxidants can induce widely different interspecific effects on lifespan.

    supplementary data available here

    pdf available from royal society pubishing website here

    and also here

  • {390}

    Shore, A.M., Karamitri, A., Kemp, P.R., SPEAKMAN, J.R., Graham, N.S and Lomax, M.A. (2013)

    Cold induced changes in gene expression in brown adipose tissue, white adipose tissue and liver.

    PLoS ONE 8: e68933


    Cold exposure imposes a metabolic challenge to mammals that is met by a coordinated response in different tissues to
    prevent hypothermia. This study reports a transcriptomic analysis in brown adipose tissue (BAT), white adipose (WAT) and
    liver of mice in response to 24 h cold exposure at 8uC. Expression of 1895 genes were significantly (P,0.05) up- or downregulated
    more than two fold by cold exposure in all tissues but only 5 of these genes were shared by all three tissues, and
    only 19, 14 and 134 genes were common between WAT and BAT, WAT and liver, and BAT and liver, respectively. We
    confirmed using qRT-PCR, the increased expression of a number of characteristic BAT genes during cold exposure. In both
    BAT and the liver, the most common direction of change in gene expression was suppression (496 genes in BAT and 590
    genes in liver). Gene ontology analysis revealed for the first time significant (P,0.05) down regulation in response to cold, of
    genes involved in oxidoreductase activity, lipid metabolic processes and protease inhibitor activity, in both BAT and liver,
    but not WAT. The results reveal an unexpected importance of down regulation of cytochrome P450 gene expression and
    apolipoprotein, in both BAT and liver, but not WAT, in response to cold exposure. Pathway analysis suggests a model in
    which down regulation of the nuclear transcription factors HNF4a and PPARa in both BAT and liver may orchestrate the
    down regulation of genes involved in lipoprotein and steroid metabolism as well as Phase I enzymes belonging to the
    cytochrome P450 group in response to cold stress in mice. We propose that the response to cold stress involves decreased
    gene expression in a range of cellular processes in order to maximise pathways involved in heat production.

  • {389}

    SPEAKMAN, J.R. (2013)

    Functional analysis of seven genes linked to body mass index (BMI) and adiposity by genome wide association studies (GWAS): a review.

    Human heredity 75: 57-79

    Brief summary: This paper reviews the potential mechanisms by which seven potential obesity related genes identified by GWAS may exert their effects on energy balance and fat storage.

    Chinese summary: 这篇论文主要综述了通过全基因组关联分析确定的7个和肥胖相关的基因在能量平衡和脂肪储存方面发挥作用的潜在机制。


    Genome-wide association studies (GWAS) have identified a
    total of about 40 single nucleotide polymorphisms (SNPs)
    that show significant linkage to body mass index, a widely
    utilised surrogate measure of adiposity. However, only 8 of
    these associations have been confirmed by follow-up GWAS
    using more sophisticated measures of adiposity (computed
    tomography). Among these 8, there is a SNP close to the
    gene FTO which has been the subject of considerable work
    to diagnose its function. The remaining 7 SNPs are adjacent
    to, or within, the genes NEGR1, TMEM18, ETV5, FLJ35779,
    LINGO2, SH2B1 and GIPR , most of which are less well studied
    than FTO, particularly in the context of obesity. This article
    reviews the available data on the functions of these genes,
    including information gleaned from studies in humans and
    animal models. At present, we have virtually no information
    on the putative mechanism associating the genes FLJ35779
    and LINGO2 to obesity. All of these genes are expressed in
    the brain, and for 2 of them ( SH2B1 and GIPR ), a direct link to

    the appetite regulation system is known. SH2B1 is an enhancer
    of intracellular signalling in the JAK-STAT pathway,
    and GIPR is the receptor for an appetite-linked hormone
    (GIP) produced by the alimentary tract. NEGR1, ETV5 and
    SH2B1 all have suggested roles in neurite outgrowth, and
    hence SNPs adjacent to these genes may affect development
    of the energy balance circuitry. Although the genes
    have central patterns of gene expression, implying a central
    neuronal connection to energy balance, for at least 4 of them
    (NEGR1, TMEM18, SH2B1 and GIPR) , there are also significant
    peripheral functions related to adipose tissue biology. These
    functions may contribute to their effects on the obese phenotype.

    pdf available at Karger web site

    and here

  • {388}

    Yang, D.B., Li, L., Wang, L.P., Chi, Q.S., Hambly, C., Wang, D.H. and SPEAKMAN, J.R. (2013)

    Limits to sustained energy intake XIX: A test of the heat dissipation limitation hypothesis in Mongolian gerbils (Meriones unguiculatus)

    Journal of Experimental Biology 216: 3358-3368

    Brief summary: During lactation mongolian gerbils appear to be limited by their capacity to dissipate heat at ambient temperatures above 21 oC, but at lower ambient temperatures are limited by some other unresolved factor. Lactating gerbils thinned their pelages to assist in heat dissipation.

    Chinese summary: 哺乳期的蒙古沙鼠在周围环境温度高于21度时会限制他们热耗散的能力,而在较低温度下的限制是由于其他一些没有被解决的因素所导致,哺乳期的沙鼠通过脱毛来辅助热耗散。


    We evaluated factors limiting lactating Mongolian gerbils (Meriones unguiculatus) at three temperatures (10oC, 21oC and 30oC). Energy intake and daily energy expenditure (DEE) increased with decreased ambient temperature. At peak lactation (day 14 of lactation), energy intake increased from 148.7 ± 5.7 at 30oC to 213.1 ± 8.2 at 21oC and 248.7 ± 12.3 at 10oC. DEE increased from 105.1 ± 4.0 at 30oC to 134.7 ± 5.6 at 21oC and 179.5 ± at 10oC on days 14-16 of lactation. With nearly identical mean litter sizes lactating gerbils at 30oC exported 32.0 less energy as milk at peak lactation, than those allocated to 10oC or 21oC, with no difference between the latter groups. On day 14 of lactation, the litter masses at 10oC and 30oC were 12.2 g and 9.3 g lower than those at 21oC, respectively. Lactating gerbils had higher thermal conductance of the fur, and lower UCP-1 levels in brown adipose tissue than non-reproductive gerbils, independent of ambient temperature, suggesting they were attempting to avoid heat stress. Thermal conductance of the fur was positively related to circulating prolactin levels. We implanted non-reproductive gerbils with mini-osmotic pumps that delivered either prolactin or saline. Prolactin did not influence thermal conductance of the fur, but did reduce physical activity and UCP-1 levels in BAT. Transferring lactating gerbils from warm to hot conditions resulted in reduced milk production, consistent with the heat dissipation limit theory, but transferring them from warm to cold conditions did not elevate milk production, consistent with peripheral limitation hypothesis or constraints on pup growth.

  • {387}

    Gibson, A.R., Ojiambo, R., Konstabel, K., Lieberman, D.E., Reilly, J.J., SPEAKMAN, J.R. and Pitsiladis, Y.P. (2013)

    Aerobic capacity, activity levels and daily energy expenditure in male and female adolescents of the Kenyan Nandi sub-group.

    PLoS ONE 8 e66552


    The relative importance of genetic and socio-cultural influences contributing to the success of east Africans in enduranceathletics remains unknown in part because the pre-training phenotype of this population remains incompletely assessed. Here cardiopulmonary fitness, physical activity levels, distance travelled to school and daily energy expenditure in 15 habitually active male (13.961.6 years) and 15 habitually active female (13.961.2) adolescents from a rural Nandi primary school are assessed. Aerobic capacity ( _VVO2max) was evaluated during two maximal discontinuous incremental exercise tests; physical activity using accelerometry combined with a global positioning system; and energy expenditure using the doubly labelled water method. The _VVO2max of the male and female adolescents were 73.965.7 ml. kg21. min21 and 61.566.3 ml. kg21. min21, respectively. Total time spent in sedentary, light, moderate and vigorous physical activities per day was 406663 min (50% of total monitored time), 244656 min (30%), 75618 min (9%) and 82630 min (10%). Average total daily distance travelled to and from school was 7.563.0 km (0.8–13.4 km). Mean daily energy expenditure, activityinduced energy expenditure and physical activity level was 12.263.4 MJ. day21, 5.463.0 MJ. day21 and 2.260.6. 70.6% of the variation in _VVO2max was explained by sex (partial R2 = 54.7%) and body mass index (partial R2 = 15.9%). Energy expenditure and physical activity variables did not predict variation in _VVO2max once sex had been accounted for. The highly active and energy-demanding lifestyle of rural Kenyan adolescents may account for their exceptional aerobic fitness and collectively prime them for later training and athletic success.

  • {386}

    Skibiel, A.L., SPEAKMAN, J.R. and Hood, W.R. (2013)

    Testing the prediction of energy allocation decisions in the evolution of life history trade-offs.

    Functional Ecology


    Brief summary: experimental manipulations of litter size increased energy demands placed on mothers but had no negative consquences for their survival or future reproductive success. However pups from enlarged litters had lower overwinter survival.

    Chinese summary:胎仔数的调整将会增加母亲能量的需求但是对他们存活率和未来交配的成功率没有消极的影响。但是来自扩大的胎仔数的窝的小崽的过冬存活率会降低。


    1. Allocating a greater amount of limited resources, such as energy, to current reproduction can reduce the amount of energy available for somatic maintenance and can ultimately impair future breeding success or maternal survival (i.e. cost of reproduction hypothesis). Although there is some support for the cost of reproduction hypothesis in birds, few empirical studies of mammals have demonstrated a trade-off between current and future reproduction. More importantly, most studies testing ultimate costs have neglected to confirm that the proximate costs of reproduction are high.
    2. We experimentally manipulated litter size in a wild population of Columbian ground squirrels for 2 years to examine both the proximate energetic and ultimate fitness (i.e. survival and breeding) costs of reproduction. We predicted that females raising augmented litters would have the highest rates of daily energy expenditure and as a result would experience lower survival rates or future fecundity.
    3. Females raising augmented litters weaned more pups, had the highest litter masses at weaning, and had field metabolic rates that were almost 1.5 times greater than females raising control or reduced litters. Contrary to our prediction there were no negative impacts of greater maternal investment and higher energy expenditure on the probability of maternal survival or future reproduction.
    4. Pups from augmented litters grew more slowly during the lactation period, were smaller at weaning and had a lower probability of survival over-winter. Thus, although females were capable of raising more young than they gave birth to without short-term costs of reduced survival or fecundity, our observations suggest that limitations to litter size are not due to a tradeoff in the allocation of energy, but rather due to the reduced survival of offspring from larger litters.
    5. Examining the proximate mechanisms hypothesized to underlie life history tradeoffs can be challenging but is critical for a comprehensive understanding of the evolution of life histories.

    pdf available at wiley website here

    lay summary also available here

  • {385}

    Wood, K.A., Stillman, R.A., Wheeler, D., Groves, S., Hambly, C., SPEAKMAN, J.R., Daunt, F. and O’Hare, M.T. (2013)

    Go with the flow: water velocity regulates herbivore foraging decisions in river catchments.

    Oikos 122: 1720-1729



    Foragers typically attempt to consume food resources that offer the greatest energy gain for the least cost, switching between habitats as the most profitable food resource changes over time. Optimal foraging models require accurate data on the gains and costs associated with each food resource to successfully predict temporal shifts. Whilst previous studies have shown that seasonal changes in food quantity and quality can drive habitat shifts, few studies have shown the effects on habitat choice of seasonal changes in metabolic foraging costs. In this study we combined field and literature data to construct an optimal foraging model to examine the effect of seasonal changes in food quantity, food quality and foraging costs on the timing of a switch from terrestrial to aquatic habitat by non-breeding mute swans Cygnus olor in a shallow river catchment. Feeding experiments were used to quantify the functional response of swans to changes in aquatic plant biomasses. By sequentially testing alternative models with fixed or variable values for food quantity, food quality and foraging cost, we found that we needed to include seasonal variance in foraging costs in the model to accurately predict the observed habitat switch date. However, we did not need to include seasonal variance in food quantity and food quality, as accurate predictions could be obtained with fixed values for these two parameters. Therefore, the seasonal changes in foraging costs were the key factor influencing the behavioural decision to switch feeding habitats. These seasonal changes in foraging costs were driven by changes in water velocity; the profitability of aquatic foraging was negatively related to water velocity, as faster water required more energy to be expended in swimming. Our results demonstrate the importance of incorporating seasonal variation in foraging costs into our understanding of the foraging decisions of animals.

    pdf available on wiley website

  • {384}

    Elliott, K.H., Ricklefs, R.E., Gaston A.J., Hatch, S., SPEAKMAN, J.R. and Davoren, G.K. (2013)

    High flight costs, but low dive costs, in auks support the biomechanical hypothesis for flightlessness in penguins s

    Proceedings of the National Academy of Science 110:9380-9384

    Data archived at data dryad.com

    media summary and impact word doc

    Brief summary : Measurements of energy demands of diving and flying in murres reveal the highest costs of flight ever measured combined with extremely efficient diving capabilities. This combination is consistent with the idea that penguins lost the abilit to fly because it is impossible to design a wing that is both good for flying and for diving.

    Chinese summary:对海鸭子在潜水和飞行的能量需求的测量揭示了在最高的飞行成本和极高潜水能力的效率相结合。这种结合的结论和企鹅失去了飞行能力因为进化出一对翅膀既擅长飞行又擅长潜水是不可能的是相一致的。


    Flight is a key adaptive trait. Despite its advantages, flight has been lost in several groups of birds, notably among seabirds, where flightlessness has evolved independently in at least five lineages. One hypothesis for the loss of flight among seabirds is that animals moving between different media face tradeoffs between maximizing function in one medium relative to the other. In particular, biomechanical models of energy costs during flying and diving suggest that a wing designed for optimal diving performance should lead to enormous energy costs when flying in air. Costs of flying and diving have been measured in free-living animals that use their wings to fly or to propel their dives, but not both. Animals that both fly and dive might approach the functional boundary between flight and nonflight. We show that flight costs for thickbilled murres (Uria lomvia), which are wing-propelled divers, and pelagic cormorants (Phalacrocorax pelagicus) (foot-propelled divers), are the highest recorded for vertebrates. Dive costs are high for cormorants and low for murres, but the latter are still higher than for flightless wing-propelled diving birds (penguins). For murres, flight costs were higher than predicted from biomechanical modeling, and the oxygen consumption rate during dives decreased with depth at a faster rate than estimated biomechanical costs. These results strongly support the hypothesis that function constrains form in diving birds, and that optimizing wing shape and form for wing-propelled diving leads to such high flight costs that flying ceases to be an option in larger wing-propelled diving seabirds, including penguins.

  • {383}

    Zhan, Y.Z., Guo, X.G., SPEAKMAN, J.R., Zuo, X.H., Wu, D., Wang, Q.H. and Yang, Z.H. (2013).

    Abundances and host relationships of Chigger mites in Yunnan Province, China.

    Medical and Veterinary entomology 27: 194-202

    Brief summary: reports details of over 92,000 chigger mites of 224 species collected from over 10,000 small mammals of 62 species living in Yunnan, China.

    Chinese summary: 本文主要报到了从栖息在中国云南的超过10,000只的62种小型哺乳动物身上收集到的224种超过92,000只恙螨的细节。


    This paper reports on ectoparasitic chigger mites found on small mammals in Yunnan Province, southwest China. Data were accumulated from 19 investigation sites (counties) between 2001 and 2009. A total of 10 222 small mammal hosts were captured and identified; these represented 62 species, 34 genera and 11 families in five orders. From the body surfaces of these 10 222 hosts, a total of 92 990 chigger mites were collected and identified microscopically. These represented 224 species, 22 genera and three subfamilies in the family Trombiculidae (Trombidiformes). Small mammals were commonly found to be infested by chigger mites and most host species harboured several species of mite. The species diversity of chigger mites in Yunnan was much higher than diversities reported previously in other provinces of China and in other countries. A single species of rodent, Eothenomys miletus (Rodentia: Cricetidae), carried 111 species of chigger mite, thus demonstrating the highest species diversity and heaviest mite infestation of all recorded hosts. This diversity is exceptional compared with that of other ectoparasites. Of the total 224 mite species, 21 species accounted for 82.2% of all mites counted. Two species acting as major vectors for scrub typhus (tsutsugamushi disease), Leptotrombidium scutellare and Leptotrombidium deliense, were identified as the dominant mite species in this sample. In addition to these two major vectors, 12 potential or suspected vector species were found. Most species of chigger mite had a wide range of hosts and low host specificity. For example, L. scutellare parasitized 30 species of host. The low host specificity of chigger mites may increase their probability of encountering humans, as well as their transmission of scrub typhus among different hosts. Hierarchical clustering analysis showed that similarities between different chigger mite communities on the 18 main species of small mammal host did not accord with the taxonomic affinity of the hosts. This suggests that the distribution of chigger mites may be strongly influenced by the environment in which hosts live.

  • {382}

    Huang, L.Q., Guo, X.G., SPEAKMAN J.R., and Dong W.G. (2013)

    Analysis of gamsid mites (Acari:Mesostigmata) associated with the Asian house rat Rattus tanezumi (Rodentia:Muridae) in Yunnan Province, Southwest China.

    Parasitology research 112: 1967-1972

    Brief summary: reports the mite populations found on over 4000 asian house rats captured over an18 year period in Yunnan, SW China. In total over 19000 mites of 50 different species were recorded.

    Chinese summary: 本文主要报道了在中国西南地区云南超过18年的时间里捕获的超过4000只亚洲家鼠身上的螨种群,有超过50种19,000只的螨的信息的记录信息。


    During a survey lasting from 1990 to 2008, we captured 4,113 Asian house rats, Rattus tanezumi Temminck 1844 (Rodentia: Muridae) from 28 counties of Yunnan Province in Southwestern China. From these rats, a total of 19,304 gamasid mites (Acari: Mesostigmata) were collected and identified as comprising 50 different species. The species diversity of gamasid mites from this single rat species is higher than that reported previously from multiple hosts within a given geographical region. Of the 50 mite species, 31 species belonged to ectoparasites and 19 species belonged to free-living mites. The species diversity of the mites from rats trapped outdoors was much higher than from rats trapped indoors. The parameter K from the negative binomial distribution was used to measure the spatial distribution patterns of the dominant mite species and revealed that all the mites had an aggregated distribution among the rat hosts. Most mite species showed a predominantly female-biased population structure with many more females than males.

  • {381}

    Liu, X.M., Zheng, Z.J., Zhu, X.M., Meng, M.H., Li, L., Shen, Y.Y., Chi, Q.S., Wang, D.H., Zhang, Z.Y., Li, C.Z., Li, Y.M., Xue, Y.M.,SPEAKMAN, J.R., and Jin, W.Z. (2013)

    Brown adipose tissue directly regulates whole body energy metabolism and energy balance.

    Cell research 23: 851-854

    Brief summary: Transplanting brown adipose tissue into mice so that they have an increased complement of this tissue both prevents weight gain on a high fat diet and reverse obesity in high fat diet induced obese mice. implantation of other tissues does not have the same effects.

    Chinese summary: 给小鼠移植褐色脂肪组织会增加这一组织,在高脂食物的诱导下会防止脂肪的增加同时还会扭转由于高脂食物诱导的小鼠肥胖。移植其他的组织将不会有相同的效果

  • {380}

    Lutermann, H., Bennett, N.C., SPEAKMAN, J.R. and Scantlebury, D.M. (2013)

    Energetic benefits of sociality offset the costs of parasitism in a cooperative mammal

    PLOS One 8: e57969

  • {379}

    Elliott, K.H., Welcker, J., Gaston, A.J., Hatch, S.A., Palace, V., Hare, J.F., SPEAKMAN, J.R. and Anderson, W.G. (2013)

    Thyroid hormones correlate with resting metabolic rate, not daily energy expenditure, in two charadriiform seabirds.

    Biology Open 2:580-586


    Both basal and sustained metabolic rate vary considerably among individuals, and are often correlated with each other when compared across species. A potential explanation for the correlation between basal and sustained metabolic rates is that they are mechanistically linked because adjustments that maximize daily energy intake also increase basal metabolic rate. As thyroid hormones are known to increase basal metabolic rate in the lab, a mechanistic linkage between basal and sustained energy expenditure would imply a correlation of these parameters with thyroid hormones. In two free-ranging bird species with high levels of energy expenditure (the black-legged kittiwake, Rissa tridactylaand thick-billed murre, Uria lomvia) we examined the relationships between free and bound levels of the thyroid hormones thyroxine (T4) and triiodothyronine (T3) with daily energy expenditure (DEE) and with 4-hr long measurements of post-absorptive and thermoneutral resting metabolism, which approximates basal metabolic rate (BMR). BMR during incubation did not correlate with DEE during chick-rearing. Free T3 (but not free T4) increased with BMR but not DEE in both species. Free T3 and free T4 did not correlate with one another. DEE correlated with body mass in kittiwakes but not murres, presumably because the larger intersexual differences in body mass for kittiwakes relative to murres had a more pronounced impact on DEE. We suggest that BMR and DEE are not linked at the level of the individual in these species and that T3, especially free T3, is a good proxy for BMR but not DEE.

  • {378}

    Zhang, L.N., Morgan, D., Sinclair, R., Selman, C., Mitchell, S.E., Clapham, J.C. and SPEAKMAN, J.R. (2013)

    Effects of chronic oral administration of a melanin-concentrating hormone receptor 1 (MCHR1) antagonist (GW803430) on energy budgets and glucose homeostasis in diet-induced obese C57BL/6J mice

    Obesity 22: 681-690

  • {377}

    SPEAKMAN, J.R. (2013)

    Measuring energy metabolism in the mouse - theoretical, practical and analytical considerations

    Frontiers in Physiology 4: article 34


    The mouse is one of the most important model organisms for understanding human genetic function and disease. This includes characterisation of the factors that influence energy expenditure and dysregulation of energy balance leading to obesity and its sequalae. Measuring energy metabolism in the mouse presents a challenge because the animals are small, and in this respect it presents similar challenges to measuring energy demands in many other species of small mammal. This paper considers some theoretical, practical and analytical considerations to be considered when measuring energy expenditure in mice. Theoretically total daily energy expenditure is comprised of several different components: basal or resting expenditure, physical activity, thermoregulation and the thermic effect of food. Energy expenditure in mice is normally measured using open flow indirect calorimetry apparatus. Two types of system are available – one of which involves a single small Spartan chamber linked to a single analyser, which is ideal for measuring the individual components of energy demand. The other type of system involves a large chamber which mimics the home cage environment and is generally configured with several chambers per analyser. These latter systems are ideal for measuring total daily energy expenditure but at present do not allow accurate decomposition of the total expenditure into its components. The greatest analytical challenge for mouse expenditure data is how to account for body size differences between individuals. This has been a matter of some discussion for at least 120 years. The statistically most appropriate approach is to use ANCOVA with individual aspects of body composition as independent predictors.

  • {376}

    Ojiambo, R., Gibson, A.R., Konstabel, K., Lieberman, D.E., SPEAKMAN, J.R., Reilly, J.J. and Pitsiladis, Y.P. (2013)

    Free-living energy expenditure of rural children and adolescents from Nandi, Kenya

    Annals of Human Biology 40: 318-323.


    PURPOSE: To examine the relationship between physical activity and energy demands in highly active lifestyles. METHODS: Physical activity patterns in 30 active Kenyan children and adolescents (14 ± 1 yrs, mean ± SD), with median body mass index (BMI) z-score -1.06 [-3.29 - 0.67] median [range], were assessed by accelerometry over 1 week. Daily energy expenditure (DEE), activity-induced energy expenditure (AEE) and Physical activity level (PAL) were simultaneously determined using doubly labelled water (DLW). Active commuting to school was assessed by global positioning system. RESULTS: Mean DEE, AEE and PAL were 12.2 ± 3.4, 5.7 ± 3.0 MJ/d and 2.3 ± 0.6, respectively. A model combining body mass (BM), average accelerometer counts per minute (CPM) and time in light activities predicted 45% of the variance in DEE (P<0.05); with a standard error of DEE estimate of 2.7 MJ/d. Furthermore, AEE accounted for approximately 47% of DEE. Distance to school was not related to variation in DEE, AEE or PAL and there was no association between active commuting and adiposity. CONCLUSION: High physical activity was associated with much higher levels of energy expenditure than observed in western societies. This contradicts the concept of energy expenditure being stable and constrained in humans.

  • {375}

    Zhao, Z. J., Song, D. G., Su, Z. C., Wei, W. B., Liu, X. B., SPEAKMAN, J. R. (2013)

    Limits to sustained energy intake XVIII: Energy intake and reproductive output during lactation in Swiss mice raising small litters

    Journal of Experimental Biology 216: 2349-2358

    doi: 10.1242/jeb.078436

    Brief summary: Previous work indicated lactating swiss mice in the cold do not upregulate their milk production. This could be a limit in milk secretory capacity of the mammary gland or a limit in pup growth capacity. By reducing litter sizes in the cold we show that pups can grow faster, implicating the mammary gland synthetic capacity as the origin of the limit.

    Chinese summary: 之前的工作暗示哺乳期的瑞士小鼠在冷暴露种不会上调他们的乳汁产量。这可能是由于乳腺的乳汁的分泌能力以及小崽生长能力所限制的。通过在冷暴露中降低胎仔数我们发现小崽的生长更快,这表明乳腺的合成能力是这种限制能力的根源。


    Limits to sustained energy intake (SusEI) during lactation in Swiss mice have been suggested to reflect the secretory capacity of the mammary glands. However, an alternative explanation is that milk production and food intake are regulated to match limited growth capacity of the offspring. In the present study, female Swiss mice were experimentally manipulated in two ways – litter sizes were adjusted to be between 1 and 9 pups and mice were exposed either to warm (21 oC) or cold (5 oC) in late lactation. Food intake, litter size and mass, milk energy output (MEO), thermogenesis, weight of the mammary glands and brown adipose tissue cytochrome c oxidase (BAT COX) activity were measured. Pup mass at weaning was almost independent of litter size at 21 oC. Positive correlations were observed between litter size, litter mass, asymptotic food intake and MEO. These data were consistent with the hypothesis that in small litters pup growth is the major factor limiting milk production. However, on average pups raised at 5oC were smaller than those raised at 21 oC. This was despite the fact milk production and food intake at the same litter sizes were both substantially higher in females raising pups at 5 oC. This suggested pup growth capacity was lower in the cold, perhaps due to pups allocating ingested energy to fuel metabolism. Differences in observed levels of milk production in different conditions may then reflect a complex interplay between factors limiting maternal performance (peripheral limitation and heat dissipation: generally better when it is cooler) and factors influencing maximum pup growth (litter size and temperature: generally better when it is hotter), and may together result in an optimal temperature favouring reproduction.

  • {374}

    Duah, O.A., Monney, K.A., Hambly, C., Król, E., and SPEAKMAN, J.R. (2013)

    Limits to sustained energy intake XVII: Lactation performance in MF1 mice is not programmed by foetal number during pregnancy

    Journal of Experimental Biology 216: 2339-2348


    Brief summary: previous work has suggested lactation performance in mice may be primed by events in pregnancy (specifically foetus number). We manipulated litter size of mice shortly after birth and no evidence that litter size in pregnancy affected lactation performance.

    Chinese summary: 之前的工作表明小鼠的哺乳行为可能是由于怀孕期间的时间(特别是胎鼠的数目)影响的。我们在小鼠出生后很短的时间内改变胎仔的数目,没有证据表明怀孕时的胎仔数会影响哺乳行为。


    Several studies have suggested that lactation performance may be programmed by the number of foetuses during pregnancy, whereas other studies indicate that processes during lactation are more important. As gestation litter size (LS) and LS in lactation are usually strongly correlated, separating the roles of pregnancy and lactation on lactation performance is difficult. To break this link, we experimentally manipulated LS of MF1 mice to 5 or 16 pups per litter by cross-fostering. LS and mass at birth were recorded on day 1 of lactation prior to LS manipulation. Maternal body mass and food intake, LS and litter mass were measured daily throughout. After weaning, mothers were euthanised and dissected to establish the potential differential utilisation of body tissues. Relationships between maternal mass and food intake, including asymptotic daily food intake at peak lactation, offspring traits and other maternal parameters suggested that the number of foetuses the females had carried during pregnancy, had no effect on lactation performance. Litter mass increases depended only on maternal food intake, which was highly variable between individuals, but was independent of foetal LS. Sizes of key organs and tissues like the liver and alimentary tract were not related to maximal food intake at peak lactation, or to foetal LS, but the masses of the pelage, mammary glands and retroperitoneal fat pad were. These data suggested that while growth of the mammary glands and associated structures may be initiated in gestation, and vary in relation to the number of placentas, the ultimate sizes and activities of the tissues depends primarily on factors during lactation.

  • {373}

    Gamo, Y., Bernard, A., Mitchell, S.E., Hambly, C., Al Jothery, A., Vaanholt, L.M., Król, E., and SPEAKMAN, J.R. (2013)

    Limits to sustained energy intake XVI: body temperatures and physical activity of female mice during pregnancy

    Journal of Experimental Biology 216


    Although lactation is the most energy demanding phase of mammalian reproduction, lactation performance may be affected by events during pregnancy. For example food intake may be limited in late pregnancy by competition for space in the abdomen between the alimentary tract and foetuses. Hence females may need to compensate their energy budgets during pregnancy by reducing activity and lowering body temperature. We explored the relationships between energy intake, body mass, body temperature and physical activity throughout pregnancy in the MF1 mouse. Food intake and body mass of 26 females were recorded daily throughout pregnancy. Body temperature and physical activity were monitored every minute for 23 hours a day by implanted transmitters (MiniMitter Inc). Body temperature and physical activity declined as pregnancy advanced, while energy intake and body mass increased. Compared to a pre-mating baseline period, mice increased energy intake by 56% in late pregnancy. Although body temperature declined as pregnancy progressed, this served mostly to reverse an increase between baseline and early pregnancy. Reduced physical activity may compensate the energy budget of pregnant mice but body temperature changes do not. Over the last 3 days of pregnancy, food intake declined. Individual variation in energy intake in the last phase of pregnancy was positively related to litter size at birth. As there was no association between the increase in body mass and the decline in intake we suggest the decline was not caused by competition for abdominal space. These data suggest overall reproductive performance is probably not constrained by events during pregnancy.

  • {372}
    1. Zhao, Z.J., Krol., E., Moille, S., Gamo, Y. and SPEAKMAN, J.R. (2013)

    Limits to sustained energy intake XV: Effects of wheel-running on the energy budget during lactation

    Journal of Experimental Biology 216: 2316-2327

    doi: 10.1242/jeb.078402


    The capacity of animals to dissipate heat may constrain sustained energy intake during lactation. We examined these constraints at peak lactation in MF1 mice that had ad libitum access to food, or that had to run a pre-set target on running wheels to obtain ad libitum access to food. The voluntary distance run decreased sharply during pregnancy and peak lactation. When lactating females were provided with 80% of their estimated food requirements, and had to run pre-set distances of 2, 4 or 6 km before given access to additional ad libitum food, most of them did not complete the running target during late lactation and the mice with the highest targets failed to reach their targets earlier in lactation. There were consequently significant group differences in asymptotic food intake (2 km, 16.97±0.40 g/d; 4 km, 14.29±0.72 g/d and 6 km, 12.65±0.45 g/d) and weaned litter masses (2 km, 71.11±2.39 g; 4 km, 54.63±4.28 g and 6 km, 47.18±2.46 g). When the females did run sufficiently to gain ad libitum food access their intake did not differ between the different distance groups or from controls that were not required to run. Thus, despite being physically capable of running the distances, mice could not exercise sufficiently in lactation to gain regular ad libitum access to food, probably because of the risks of hyperthermia when combining heat production from exercise with thermogenesis from lactation.

  • {371}

    Vaanholt, L.M., Sinclair, R. and SPEAKMAN, J.R. (2013)

    Limits to sustained energy intake XIV: Heritability of reproductive performance in mice

    Journal of Experimental Biology 216 2308-2315

    doi: 10.1242/jeb.078394

    Brief summary: cross fostering experiments show that asymptotic food intake at peak lactation and milk production are highly heritable traits in mice.

    Chinese summary: 交叉培养实验表明小鼠哺乳和产奶高峰期渐近的食物摄入量是高度遗传的。


    Limits to sustained energy intake (SusEI) are important because they constrain many aspects of animal performance. Individual variability in SusEI may be imposed by genetic factors that are inherited from parents to offspring. Here we investigated heritability of reproductive performance in MF1 mice. Food intake, milk energy output (MEO) and litter mass was measured in mothers (F0) and daughters (F1) that were raising litters of 10 pups. Cross fostering was designed so that half of each litter consisted of biological offspring and the rest came from one unrelated female (i.e., fostered pups). Food intake increased linearly during early lactation and reached a plateau during late lactation (day 9-13, called the asymptotic food intake; FIAS equivalent to SusEI). Parent-offspring regression showed that FIAS, MEO and litter mass were all positively and significantly related between mothers and their biological daughters, but no significant relationships were found between the same traits for mothers and fostered daughters. Asymptotic food intake at peak lactation was significantly correlated to adult food intake and body mass when the mice were 6 months old and not lactating. In conclusion, a large part of the variation in FIAS could be explained by genetic variation or maternal effects in pregnancy whereas non-genetic maternal effects in lactation were negligible. As a consequence, biological daughters of mothers with high reproductive performance (i.e., high milk production and hence higher litter mass at weaning) had a better reproductive performance themselves, independent of the mother that raised them in lactation.

  • {370}
    1. Welcker,J., Chastel, O., Gabrielsen, G.W., Guillaumin, J., Kitaysky, A.S., SPEAKMAN, J.R., Tremblay, Y. and Bech, C. (2013)

    Thyroid hormones correlate with basal metabolic rate but not field metabolic rate in free-living kittiwakes

    PLOS One 8: e56229


    Thyroid hormones (TH) are known to stimulate in vitro oxygen consumption of tissues in mammals and birds. Hence, in
    many laboratory studies a positive relationship between TH concentrations and basal metabolic rate (BMR) has been
    demonstrated whereas evidence from species in the wild is scarce. Even though basal and field metabolic rates (FMR) are
    often thought to be intrinsically linked it is still unknown whether a relationship between TH and FMR exists. Here we
    determine the relationship between the primary thyroid hormone triiodothyronine (T3) with both BMR and FMR in a wild
    bird species, the black-legged kittiwake (Rissa tridactyla). As predicted we found a strong and positive relationship between
    plasma concentrations of T3 and both BMR and mass-independent BMR with coefficients of determination ranging from
    0.36 to 0.60. In contrast there was no association of T3 levels with either whole-body or mass-independent FMR (R2 = 0.06
    and 0.02, respectively). In accordance with in vitro studies our data suggests that TH play an important role in modulating
    BMR and may serve as a proxy for basal metabolism in wild birds. However, the lack of a relationship between TH and FMR
    indicates that levels of physical activity in kittiwakes are largely independent of TH concentrations and support recent
    studies that cast doubt on a direct linkage between BMR and FMR.

  • {369}

    Archibald, D.W., Fletcher, Q.E., Boutin, S., McAdam, A. G., SPEAKMAN, J.R. and Humphries, M.M. (2013)

    Sex-specific hoarding behavior in North American red squirrels (Tamiasciurus hudsonicus)

    Journal of Mammalogy 94: 761-770.

  • {368}
    • SPEAKMAN, J.R. (2013)

      Evolutionary perspectives on the obesity epidemic: adaptive, maladaptive and neutral viewpoints

      Annual review of nutrition 33: 289-317


      The prevalence of obesity in modern societies has two major contributory factors---an environmental change that has happened in historical times and a genetic predisposition that has its origins in our evolutionary history. Understanding both aspects is complex. From an evolutionary perspective, three different types of explanation have been proposed. The first is that obesity was once adaptive and enabled us to survive (or sustain fecundity) through periods of famine. People carrying so-called thrifty genes that enabled the efficient storage of energy as fat between famines would be at a selective advantage. In the modern world, however, people who have inherited these genes deposit fat in preparation for a famine that never comes, and the result is widespread obesity. The key problem with this, and any other adaptive scenario, is to understand why, if obesity was historically so advantageous, many people did not inherit these thrifty genes and in modern society are able to remain slim, despite the environmental change favoring fat storage. The second type of explanation is that obesity is not adaptive and may never even have existed in our evolutionary past, but it is favored today as a maladaptive by-product of positive selection on some other trait. An example of this type of explanation is the suggestion that obesity results from variation in brown adipose tissue thermogenesis. Finally, a third class of explanation is that most mutations in the genes that predispose us to obesity are neutral and have been drifting over evolutionary time---so-called drifty genes, leading some individuals to be obesity prone and others obesity resistant. In this article, I review the current evidence for and against these three different scenarios and conclude that the thrifty gene hypothesis is untenable but the other two ideas may provide a cogent explanation of the modern obesity phenomenon.

  • {367}

    Elliott, K.H., Le Vaillant, M., Kato, A., SPEAKMAN, J.R. and Ropert-Coudert, Y. (2013)

    Accelerometry predicts daily energy expenditure in a bird with an active lifestyle

    Biology Letters 9: 20120919

  • {366}

    Bennett, K.A., Fedak, M.A., Moss, S.E.W., Pomeroy, P.P., SPEAKMAN, J.R. and Hall, A.J. (2013)

    The role of glucocorticoids in naturally fasting grey seal (Halichoerusgrypus) pups: dexamethasone stimulates mass loss and protein utilisation, but not departure from the colony

    Journal of Experimental Biology 216: 984-991

  • {365}
    • SPEAKMAN, J.R., Fletcher, Q.E. and Vaanholt, L.M. (2013)

      The '39 steps': an algorithm for performing statistical analysis of energy intake and expenditure data.

      Disease models and mechanisms 6: 293-301


      Brief summary: a step by step guide to the statistical analysis and interpretation of energy expenditure and energy intake data.

      Chinese summary: 统计分析分部指南和能量支出与能量摄入数据的解读


      The epidemics of obesity and diabetes have aroused great interest in the analysis of energy balance, with the use of
      organisms ranging from nematode worms to humans. Although generating energy-intake or -expenditure data is relatively
      straightforward, the most appropriate way to analyse the data has been an issue of contention for many decades. In the
      last few years, a consensus has been reached regarding the best methods for analysing such data. To facilitate using
      these best-practice methods, we present here an algorithm that provides a step-by-step guide for analysing energy -
      intake or -expenditure data. The algorithm can be used to analyse data from either humans or experimental animals,
      such as small mammals or invertebrates. It can be used in combination with any commercial statistics package; however,
      to assist with analysis, we have included detailed instructions for performing each step for three popular statistics packages
      (SPSS, MINITAB and R). We also provide interpretations of the results obtained at each step. We hope that this algorithm
      will assist in the statistically appropriate analysis of such data, a field in which there has been much confusion and some

      if you require individual help with any data set feel free to email me (

      download step by step flowchart from paper here. This is best printed off at A3 size.

      supplementary material containing details of the exact commands for each of the 39 steps for 3 common statistical packages (SPSS, MINITAB and R) is available here

      paper pdf is available from the DMM web site here

      and is also available here

  • {364}
    • Fletcher, Q.E., Selman, C., Boutin, S., McAdam, A.G., Woods, S.B., Seo, A.Y., Leeuwenburgh, C. SPEAKMAN, J.R. and Humphries, M.M. (2013)

      Oxidative damage increases with reproductive energy expenditure and is reduced by food supplementation.

      Evolution 67: 1527-1536

      doi: 10.1111/evo.12014


      A central principle in life-history theory is that reproductive effort negatively affects survival. Costs of reproduction are thought to be physiologically based, but the underlying mechanisms remain poorly understood. Using female North American red squirrels (Tamiasciurus hudsonicus), we test the hypothesis that energetic investment in reproduction overwhelms investment in antioxidant protection, leading to oxidative damage. In support of this hypothesis we found that the highest levels of plasma protein oxidative damage in squirrels occurred during the energetically demanding period of lactation. Moreover, plasma protein oxidative damage was also elevated in squirrels that expended the most energy and had the lowest antioxidant protection. Finally, we found that squirrels that were food-supplemented during lactation and winter had increased antioxidant protection and reduced plasma protein oxidative damage providing the first experimental evidence in the wild that access to abundant resources can reduce this physiological cost.

      pdf available from Wiley website

  • {363}

    Muller, T.D., Lee, S.J., Jastroch, M., Stemmer, K., Aichler, M., Abplanalp, B., Anathakrishna, G., Bhardwaj, N., Collins, S., Divanovic, S., Finan, B., Gao, Y, Habegger, K., Menbree, J., Heppner, K., Hofmann, S., Holland, J., Kutschke, M., Krishna, R., Oelkrug, R., Ottaway, N., Perez-Tilve, D., Raver, C., Walch, A.K., SPEAKMAN, J.R., Tseng, Y.H., Kahn, C.R., Spiegelman, B., Diaz-Meco, M., Pfluger, P.T., Moscat, J. and Tschop, M.H. (2013)

    p62 links beta-adrenergic input with mitochondrial function

    Journal of Clinical Investigation 123: 469-478.

  • {362}
    • McNay, D. and SPEAKMAN, J.R. (2013)

      High fat diet causes rebound weight gain

      Molecular metabolism 2:103-108

      media summary and impact word doc

      See also commentary on this article that was published in molecular metabolism to coincide with the appearance of the actual article in print.

      M. Rosenbaum, Y. Ravussin and R.L. Leibel (2013) Does diet have structural consequences in the hypothalamus? Molecular metabolism 2: 58-59.

  • {361}

    Berman, E., Fortson, S., Snaith, S., Gupta, M., Chery, I., Blanc, S., Melanson, E., Thomson, P., and SPEAKMAN, J.R. (2013)

    Direct analysis of δ2H and δ18O in natural and enriched human urine using laser-based, Off-Axis Integrated Cavity Output Spectroscopy.

    Analytical Chemistry 84: 9768-9773.

  • {360}
    • SPEAKMAN, J.R. and Keijer, J. (2013)

      Not so hot: optimal temperatures for housing mice to mimic the thermal environment of humans.

      Molecular metabolism 2: 5-9

      See also comment on this paper

      Gaskill, B.N. and Garner, J.P. (2013) Letter to the editor on “Not so hot: optimal housing temperatures for mice to mimic the thermal environment of humans. Molecular metabolism

      And our reply

      Speakman, J.R. and Keijer, J. (2013) Not so nuanced: reply to the comments by Gaskill and Garner (2013) on “Not so hot: optimal housing temperatures for mice to mimic the thermal environment of humans. Molecular metabolism

  • {359}

    SPEAKMAN, J.R. (2013)

    Sex and age related mortality profiles during famine: testing the ‘body fat’ hypothesis.

    Journal of Biosocial Science 45: 823-840

  • {358}

    Careau, V., Réale, D., Garant, D., Pelletier, F., SPEAKMAN, J.R. and Humphries, M.M. (2013)

    Context dependent correlation between resting metabolic rate and daily energy expenditure in wild chipmunks.

    Journal of Experimental Biology 216: 418-426.

  • {357}

    SPEAKMAN, J.R. and Westerterp, K.R. (2013)

    A mathematical model of weight loss under total starvation: evidence against the thrifty-gene hypothesis

    Disease Models and Mechanisms 6: 236-251

  • {356}

    Polyviou, T.P., Easton, C., Beis, L., Malkova, D., Takas, P., Hambly, C., SPEAKMAN, J.R. Koehler, K., and Pitsiladis, Y.P. (2012)

    Effects of glycerol and creatine hyperhydration on doping-relevant blood parameters

    Nutrients 4: 1171-1186.

  • {355}
    • Selman, C., Blount, J.D., Nussey, D.H. and SPEAKMAN, J.R. (2012)

      Oxidative damage, Ageing and life-history evolution - where now?

      Trends in Ecology and Evolution 27:


      The idea that resources are limited and animals can maximise fitness by trading costly activities off against one another forms the basis of life-history theory. Although investment in reproduction or growth negatively affects survival, the mechanisms underlying such tradeoffs remain obscure. One plausible mechanism is oxidative damage to proteins, lipids, and nucleic acids caused by reactive oxygen species (ROS). Here, we critically evaluate the premise that ROS-induced oxidative damage shapes life history, focussing on birds and mammals, and highlight the importance of ecological studies examining free-living animals within this experimental framework. We conclude by emphasising the value of using multiple assays to determine oxidative protection and damage. We also highlight the importance of using standardised and appropriate protocols, and discuss future research directions.

  • {354}

    Polyviou, T.P., Pitsiladis, Y.P., Lee, W.C., Takas, P., Hambly, C., SPEAKMAN, J.R., and Malkova, D. (2012)

    Thermoregulatory and Cardiovascular Responses to Creatine, Glycerol and Alpha Lipoic Acid in Trained Cyclists

    Journal of the International Society of Sports Nutrition 9: 29

  • {353}

    Careau, V., Bergeron, P., Garant, D., Réale, SPEAKMAN, J.R. and Humphries, M.M. (2013)

    The energetic and survival costs of growth in free-ranging chipmunks

    Oecologia 171: 11-23.

  • {352}

    Heitmann, B.L., Westerterp, K.R., Loos, R.J.F., Sørensen, T.I.A., O’Dea, K., McLean, P.,Jensen, T.K., Eisenmann, J., SPEAKMAN, J.R., Simpson, S.J., Reed, D.R. and Westerterp-Plantenga, M.S. (2012)

    Obesity: lessons from evolution and the environment.

    Obesity reviews 13: 910-922

  • {351}

    Gremillet, D., Welcker, J., Karnovsky, N.J., Wojcieh, W., Hall, M.E., Fort, J., SPEAKMAN, J.R. and Harding, A.M.A. (2012).

    Little auks buffer the impact of current Arctic climate change.

    Marine Ecology Progress Series.454: 197-206

  • {350}

    Fletcher, Q., SPEAKMAN, J.R., Boutin, S., McAdam, A., Woods, S. and Humphries, M.M. (2012)

    Seasonal stage differences overwhelm environmental and individual factors as determinants of energy expenditure in free-ranging red squirrels

    Functional ecology 26: 677-687

  • {349}

    Careau, V., Reale, D., Garant, D., SPEAKMAN, J.R. and Humphries, M.M. (2012)

    Stress induced rise in body temperature is repeatable in free-ranging Eastern chipmunks (Tamias striatus)

    Journal of Comparative Physiology 182: 403-414

  • {348}

    Zhang, L.N., Gamo, Y., Sinclair, R., Mitchell, S.E., Morgan, D., Clapham, J.C. and SPEAKMAN, J.R. (2012)

    Effects of chronic oral rimonabant administration on energy budgets of diet-induced obese C57BL/6J mice

    Obesity 20: 954-962.

  • {347}

    Krol, E., Martin, S.A.M., Huhtaniemi, I.T., Douglas, A., and SPEAKMAN, J.R. (2011).

    Negative correlation between milk production and brown adipose tissue gene expression in lactating mice.

    Journal of Experimental Biology214: 4160-70

  • {346}

    Bennett, K.A., Moss, S.E.W., Pomeroy, P., SPEAKMAN, J.R. and Fedak M.A. (2012).

    Effects of handling regime and sex on changes in cortisol, thyroid hormones and body mass in fasting grey seal pups

    Comparative Biochemistry and Physiology161: 69-76

  • {345}

    Jacobs, S.A., Elliott, K.H., Gaston, A.J., Guigueno, M.F., Redman,P., SPEAKMAN, J.R, and Weber, J.M. (2012)

    Determining seabird body condition using non-lethal measures: lipid dynamics at high and low arctic colonies

    Physiological and Biochemical Zoology 85: 85-95

  • {344}

    Zhang, L.N., Mitchell, S.E., Hambly, C., Morgan, D., Clapham J.C., and SPEAKMAN, J.R. (2012)

    Physiological and behavioral responses to intermittent starvation in C57BL/6J mice

    Physiology and Behaviour 105: 376-387

  • {343}

    Meakin, P.J., Harper, A.J., Hamilton, D.L., Gallagher, J., McNeilly, A.D., Burgess, L.A., Vaanholt, L.M., Bannon, K.A., Latcham, J., Hussain, I., SPEAKMAN, J.R., Howlett, D.R., and Ashford, M.L.J (2012)

    Reduction in BACE1 decreases body weight, protects against diet-induced obesity and enhances insulin sensitivity in mice.

    Biochemical Journal 441: 285-296

  • {342}

    Vaanholt, L.M., Magee, V., and SPEAKMAN, J.R. (2012)

    Factors predicting individual variability in diet-induced weight loss in MF1 mice.

    Obesity20: 285-294.

  • {341}

    Tschöp, M.H*., SPEAKMAN, J.R.*,Arch, J.R.S., Auwerx, J., Brüning, J.C., Chan, L., Eckel, R., Farese R.V., Galgani, J.E., Hambly, C., Horvath, T.L., Kahn, B.B., Kozma, S.C., Maratos-Flier, E., Müller, T.D., Muenzberg, H., Pfluger, P.T., Plum, L., Reitman, M., Rahmouni, K., Shulman, G.I., Thomas, G., Kahn, C.R., and Ravussin, E. (2012)

    A guide to analysis of mouse energy metabolism

    Nature methods 9: 57-63
    *Joint first authors

    Brief summary: consensus summary of anlytical approaches to analyse data derived from energetics studies of food intake and energy expenditure

    Chinese summary: 食物摄入和能量支出的能量学研究数据分析方法的总结

    We present a consolidated view of the complexity and challenges of designing studies for measurement of energy metabolism in mouse models, including a practical guide to the assessment of energy expenditure, energy intake and body composition and statistical analysis thereof. We hope this guide will facilitate comparisons across studies and minimize spurious interpretations of data. We recommend that division of energy expenditure data by either body weight or lean body weight and that presentation of group effects as histograms should be replaced by plotting individual data and analyzing both group and body-composition effects using analysis of covariance (ANCOVA).

  • {340}

    Hambly, C., Duncan, J.S., Archer, Z.A., Moar, K.M., Mercer, J.G. and SPEAKMAN, J.R. (2012)

    Repletion of TNF-α or leptin in calorically restricted mice suppresses post-restriction hyperphagia

    Disease Models and Mechanisms 5: 83-94


    The causes of post-restriction hyperphagia (PRH) represent a target for drug based therapies to prevent obesity. However, the factors causing PRH are poorly understood. We show in mice that the extent of PRH was independent of the time under restriction, but depended on its severity, suggesting that PRH was driven by signals from altered body composition. Signals related to fat mass were important drivers. Circulating levels of leptin and TNF-α were significantly depleted following caloric restriction (CR). We experimentally repleted their levels to match those of controls, and found that in both treatment groups the level of PRH was significantly blunted. These data establish a role for TNF-α and leptin in the non-pathological regulation of energy homeostasis. Signals from adipose tissue, including but not limited to leptin and TNF-α, regulate PRH and may be targets for therapies that support people engaged in CR to reduce obesity.

  • {339}

    SPEAKMAN, J.R.,Levitsky, D.A., Allison, D.B., Bray, M.S., de Castro, J.M., Clegg, D.J., Clapham, J.C., Dulloo, A.G., Gruer, L., Haw, S., Hebebrand, J., Hetherington, M.M., Higgs, S., Jebb, S.A., Loos, R.J.F., Luckman, S., Luke, A., Mohammed-Ali, V., O'Rahilly, S., Periera, M., Perusse, L., Robinson, T.N., Rolls, B., Symonds, M.E. and Westerterp-Plantenga, M.S. (2011)

    Set-points, settling points and some alternative models: theoretical options to understand how genes and environments combine to regulate body adiposity.

    Disease Models and Mechanisms 4:?733-745

    Brief summary: review of different models that have been proposed to understand body weight regulation. strengths and weaknesses of the different models are presented concluding that the dual-intervention point model provides the most cogent explanation of current data.

    Chinese summary: 本文通过对不同体重调节模型优缺点的比较,得出了双干预点模型是基于目前数据揭示体重调节最令人信服的解释。


    The close correspondence between energy intake and expenditure over prolonged time periods, coupled with an apparent protection of the level of body adiposity in the face of perturbations of energy balance, has led to the idea that body fatness is regulated via mechanisms that control intake and energy expenditure. Two models have dominated the discussion of how this regulation might take place. The set point model is rooted in physiology, genetics and molecular biology, and suggests that there is an active feedback mechanism linking adipose tissue (stored energy) to intake and expenditure via a set point, presumably encoded in the brain. This model is consistent with many of the biological aspects of energy balance, but struggles to explain the many significant environmental and social influences on obesity, food intake and physical activity. More importantly, the set point model does not effectively explain the ‘obesity epidemic’ – the large increase in body weight and adiposity of a large proportion of individuals in many countries since the 1980s. An alternative model, called the settling point model, is based on the idea that there is passive feedback between the size of the body stores and aspects of expenditure. This model accommodates many of the social and environmental characteristics of energy balance, but struggles to explain some of the biological and genetic aspects. The shortcomings of these two models reflect their failure to address the gene-by-environment interactions that dominate the regulation of body weight. We discuss two additional models – the general intake model and the dual intervention point model – that address this issue and might offer better ways to understand how body fatness is controlled.

    Peer-recognition: This paper was reviewed at Faculty 1000 by Jana Vliet-Ostapachouk and Harold Snieder on the 12th December 2011 and graded a ?must read' (8.0). It was also rated on the 3rdJanuary 2012 by Julian Mercer at Faculty1000 Physiology and also graded a 'must read' (8.0).

  • {338}

    SPEAKMAN, J.R.and Mitchell, S.E. (2011)

    Caloric restriction

    Molecular Aspects of Medicine 32: 159-221

    Brief summary: review of caloric restriction and its impact on longevity, including impacts on health and supposed molecular mechanisms. Ends with summary of caloric restriction mimetic drugs.

    Chinese summary:本文主要综述了能量限制以及它对寿命的影响包括对于健康的影响和假设的分子机制,结尾总结了能量限制拟合药物。


    Restricting the intake of calories has been practiced as a method for increasing both the length and quality of life for over 500 years. Experimental work confirming the success of this approach in animals has accumulated over the last 100 years. Lifelong caloric restriction (CR) may extend life by up to 50% in rodents, with progressively less impact the later in life it is started. This effect is matched by profound impacts on age related diseases including reduced risk of cancer, neurodegenerative disorders, autoimmune disease, cardiovascular disease and type II diabetes mellitus. The disposable soma theory of ageing suggests that CR evolved as a somatic protection response to enable animals to survive periods of food shortage. The shutdown of reproductive function during CR is consistent with this suggestion, but other features of the phenomenon are less consistent with this theory, and some have suggested that in rodents it may be mostly an artefact of domestication. CR induces profound effects on animals at all levels from the transcriptome to whole animal physiology and behaviour. Animals under CR lose weight which is disproportionately contributed to by white adipose tissue. Generally animals on CR change their activity patterns so that they are more active prior to food delivery each day but total activity may be unchanged or reduced. Considerable debate has occurred over the effects of CR on resting metabolic rate (RMR). Total RMR declines, but as body mass and body composition also change it is unclear whether metabolism at the tissue level also declines, is unchanged or even increases. Body temperature universally decreases. Hunger is increased and does not seem to abate even with very long term restriction. Circulating adipokines are reduced reflecting the reduction in white adipose tissue (WAT) mass under restriction and there is a large reduction in circulating insulin and glucose levels. There are profound tissue level changes in metabolism with a generalised shift from carbohydrate to fat metabolism. Four pathways have been implicated in mediating the CR effect. These are the insulin like growth factor (IGF-1)/insulin signalling pathway, the sirtuin pathway, the adenosine monophosphate (AMP) activated protein kinase (AMPK) pathway and the target of rapamycin (TOR) pathway. These different pathways may interact and may all play important roles mediating different aspects of the response. Exactly how they generate the health benefits remains open for debate, however CR results in reduced oxidative stress and enhanced autophagy, both of which could be essential components of the beneficial effects. Most data about the effects of CR in mammals comes from work on rodents. There is limited work on non-human primates that shows promising effects and one randomised controlled trial in humans where physiological markers of the CR response are consistent with the responses in mice and rats. There are also populations of humans voluntarily restricting themselves. Humans on long term restriction report similar negative side effects to those observed in animals – perpetual hunger, reduced body temperature leading to a feeling of being cold, and diminished libido. Considerable effort has been directed in recent years to find drugs that mimic the CR response. Promising candidates are those that intersect with the critical signalling pathways identified above and include biguanides such as metformin that target the insulin signalling pathway, stilbenes (e.g. resveratrol) that affect sirtuin activity and drugs such as rapamycin that interact with mTOR signalling. Whether it will ever be possible to find drugs that capture the health benefits of CR without the negative side-effects remains unclear. Moreover, even if such drugs are developed how the current licensing system for drug use in western societies would cope with them may be a further obstacle to their use.

    This was one of the top 5 most downloaded papers from the journal web site during the first 6 months of 2013.

  • {337}

    Hall, K.H., Heymsfield, S., Kemnitz, J., Klein, S., Schoeller, D.A. and SPEAKMAN, J.R. (2012)

    Energy balance and body weight regulation: a useful concept for understanding the obesity epidemic

    American Journal of Clinical Nutrition95: 989-994

  • {336}

    Cameron, K.M., Golightly, A., Miwa, S., SPEAKMAN, J.R., Boys, R. and von Zglinicki, T. (2011)

    Gross energy metabolism in mice under late onset short term caloric restriction.

    Mechanisms of Ageing and Development 132: 202-209

  • {335}

    Garcia, C., Huffman, M.A., Shimizu, K., and SPEAKMAN, J.R. (2011)

    Energetic consequences of seasonal breeding in female Japanese macaques (Macaca fuscata)

    American Journal of Physical Anthropology146: 161-170

  • {334}

    Zhang, L.N., Morgan, D., Clapham, J.C. and SPEAKMAN, J.R. (2012)

    Factors predicting non-genetic variability in body weight gain induced by a high fat diet in inbred C57BL/6J mice

    Obesity20: 1179-88

  • {333}

    Bergeron, P., Careau, V., Humphries, M.H., Réale, D., SPEAKMAN, J.R. and Garant, D. (2011)

    The energetic and oxidative costs of reproduction in a free-ranging rodent

    Functional Ecology25: 1063-1071

  • {332}

    Rutkowska J., Koskela,E., Mappes, T. and SPEAKMAN, J.R. (2011)

    A trade-off between current and future sex allocation revealed by maternal energy budget in a small mammal

    Proc. Roy Soc B. 278: 2962-2969

  • {331}

    Ahlstrom, O., Redman, P., and SPEAKMAN J.R. (2011)

    Energy expenditure and water turnover in hunting dogs in winter conditions

    British Journal of Nutrition106: S158-161.

  • {330}

    SPEAKMAN, J.R. and Selman, C. (2011)

    The free-radical damage theory: evidence accumulates against a simple link of oxidative stress to ageing and lifespan.

    Bioessays 33: 255-259

  • {329}

    Johnson, M.A., Dwyer, J.T., Jensen, G.L., Miller, J.W., SPEAKMAN, J.R., Starke-Reed, P., Volpi, E. (2011)

    Challenges and New Opportunities for Clinical Nutrition Interventions in the Aged

    Journal of Nutrition 141: 535-541

  • {328}

    Cui, J.G., Tang, G.B., Wang, D.H and SPEAKMAN, J.R. (2011)

    Effects of leptin infusion during peak lactation on food intake, body composition, litter growth and maternal neuroendocrine status in female Brandt’s voles (Lasiopodomys brandtii)

    American Journal of Physiology 300: R447-459

  • {327}

    Cameron, K.M., Morris, P.J., Hackett, R.M. and SPEAKMAN, J.R. (2011)

    The effects of increasing water content to reduce the energy density of the diet on body mass changes following caloric restriction in domestic cats.

    Journal of Animal Physiology and Animal Nutrition 95: 399-408.

    The raw data associated with this publication is made freely available for use by other scientists in the attached excel file. If you use these data please acknowledge the source as this publication.

  • {326}

    Cameron, K.M. and SPEAKMAN, J.R. (2011)

    Reduction of dietary energy density reduces body mass regain following energy restriction in female mice.

    Journal of Nutrition 141: 182-188

  • {325}

    Hinsley, S.A., Bellamy, P.E., Rothery, P., Redman, P., Furness, L., and SPEAKMAN, J.R. (2011)

    Effects of the Doubly Labelled Water Procedure on Great Tits Parus major feeding young

    Bird Study 58: 151-159

  • {324}

    Simons, M.J.P., Reimert, I., van der Vinne, V. Hambly, C., Vaanholt, L.M., SPEAKMAN, J.R. and Gerkema, M.P. (2011)

    Ambient temperature shapes reproductive output during pregnancy and lactation in the common vole (Microtus arvalis): a test of the heat dissipation limit theory.

    Journal of Experimental Biology 214: 38-49

  • {323}

    SPEAKMAN, J.R. and Krol, E. (2011)

    Limits to sustained metabolic rate XIII: recent progress and future perspectives

    Journal of Experimental Biology 214: 230-241

  • {322}

    Zub, K., Szafranska, P., Konarzewski, M., and SPEAKMAN, J.R. (2011)

    Effect of energetic constraints on distribution and winter survival of weasel males.

    Journal of Animal Ecology 80, 259–269

  • {321}

    Bennett, K.A., McConnell, B.J., Moss, S.E.W., SPEAKMAN, J.R., Pomeroy, P.P. and Fedak, M.A. (2010)

    Effects of age and body mass on development of diving capabilities of grey seal pups: costs and benefits of the post-weaning fast

    Physiological and Biochemical Zoology 83: 911-923

  • {320}

    SPEAKMAN, J.R. and Westerterp, K.R. (2010)

    Associations between energy demands, physical activity and body composition in adult humans between 18 and 96 years of age

    American Journal of Clinical Nutrition 92: 826-834

    This paper was featured on the medical review site MDLinx

  • {319}

    Nachvak, S.M., Neyestani, T.R., Keshawarz, A.S. and SPEAKMAN, J.R. (2010)

    Low consumption of fruit and vegetables, and markers of oxidative stress in children with Down syndrome.

    Down Syndrome Research and Practice

  • {318}

    Schultner, J., Welcker, J., SPEAKMAN, J.R., Norday, E.S. and Gabrielsen, G.W. (2010).

    Application of the two-sample doubly labelled water method alters behaviour and affects estimates of energy expenditure in black-legged kittiwakes

    Journal of Experimental Biology 213: 2958-2966

    This paper was featured on the front cover of the journal issue.

  • {317}

    SPEAKMAN, J.R.and Krol, E. (2010)

    The heat dissipation limit theory and evolution of life histories in endotherms - time to dispose of the disposable soma theory?

    Integrative and Comparative Biology 50:793-807


    Brief summary: The disposable soma theory posits that ageing results because costly somatic protection mechanisms must be traded off, within an envolope of available energy supply, against reproduction. This paper builds on our previous paper (number 322) where we developed a theory that endotherms may be limited by their capacity to dissipate heat. If true this theory undermines the disposable soma theory. The current paper explores these ramifications, and proposes some tests that would separate between the alternative hypotheses.

    Chinese summary: 一次性体细胞理论假设衰老是由机体保护机制必须在足够的能量供应与保证生殖两个方面权衡。这篇文章是根据我们以前发表的(编号322)完善而来的,在这篇文章我们提出恒温动物散热能力有限。如果我们的研究是正确的,那么将推翻一次性体细胞理论。这篇文章对这些分歧进行了探索并且在提出了一些可以区分这些假说的测试。


    A major factor influencing life-history strategies of endotherms is body size. Larger endotherms live longer, develop more slowly, breed later and less frequently, and have fewer offspring per attempt at breeding. The classical evolutionary explanation for this pattern is that smaller animals experience greater extrinsic mortality, which favors early reproduction at high intensity. This leads to a short lifespan and early senescence by three suggested mechanisms. First, detrimental late-acting mutations cannot be removed because of the low force of selection upon older animals (mutation accumulation). Second, genes that promote early reproduction will be favored in small animals, even if they have later detrimental effects (antagonistic pleiotropy). Third, small animals may be forced to reduce their investment in longevity assurance mechanisms (LAMs) in favor of investment in reproduction (the disposable soma theory, DST). The DST hinges on three premises: that LAMs exist, that such LAMs are energetically expensive and that the supply of energy is limited. By contrast, the heat dissipation limit (HDL) theory provides a different conceptual perspective on the evolution of life histories in relation to body size. We suggest that rather than being limited, energy supplies in the environment are often unlimited, particularly when animals are breeding, and that animals are instead constrained by their maximum capacity to dissipate body heat, generated as a by-product of their metabolism. Because heat loss is fundamentally a surface-based phenomenon, the low surface-to-volume ratio of larger animals generates significant problems for dissipating the body heat associated with reproductive effort, which then limits their current reproductive investment. We suggest that this is the primary reason why fecundity declines as animal size increases. Because large animals are constrained by their capacity for heat dissipation, they have low reproductive rates. Consequently, only those large animals living in habitats with low extrinsic mortality could survive leading to the familiar patterns of life-history trade-offs and their links to extrinsic mortality rates. The HDL theory provides a novel mechanism underpinning the evolution of life history and ageing in endotherms, and makes a number of testable predictions that directly contrast with the predictions arising from the DST.

    Peer recognition: The paper was recommended at Faculty 1000 biology by Daniel Promislow on June 1st 2010 graded as a 'must read'. It was also recommended by Marc Mangel on August 13th 2010 graded as a 'must read'.

  • {316}

    Cameron, K.M. and SPEAKMAN, J.R. (2010)

    The extent and function of ‘food grinding’ in the laboratory mouse (Mus musculus).

    Laboratory Animals 44: 298-304

    doi: 10.1258/la.2010.010002

    The raw data associated with this publication is made freely available for use by other scientists in the attached excel file. If you use these data please acknowledge the source as this publication.

  • {315}

    Wyse, C.A., Coogan, A.N., Selman, C., Hazlerigg, D.G. and SPEAKMAN, J.R. (2010)

    Association between Mammalian Lifespan and Circadian Free-Running Period: The Circadian Resonance Hypothesis Revisited

    Biology Letters 6: 696-698

    The raw data used in the analyses in this paper is made freely available for use by other scientists in the attached pdf file. If you use these data please acknowledge the source as this publication.

  • {314}

    SPEAKMAN, J.R.and Krol, E (2010)

    Maximal heat dissipation capacity and hyperthermia risk: neglected key factors in the ecology of endotherms

    Journal of Animal Ecology 79:726-746

    doi: 10.1111/j.1365-2656.2010.01689.x

    Brief summary: This paper summarises our novel theory that in many cases endothermic animals are not limited by external energy supplies or their abilities to take in energy, but rather are limited by their capacity to dissipate body heat and hence the risk of hyperthermia.

    Chinese summary:这篇文章总结了我们的新理论即在很多情况下恒温动物是由于他们自身散热能力的限制有高热的风险而不是被外界的能量供给以及自身的能量的能量摄入所限。


    1. The role of energy in ecological processes has hitherto been considered primarily from the standpoint that energy supply is limited. That is, traditional resource-based ecological and evolutionary theories and the recent ‘metabolic theory of ecology’ (MTE) all assume that energetic constraints operate on the supply side of the energy balance equation.

    2. For endothermic animals, we provide evidence suggesting that an upper boundary on total energy expenditure is imposed by the maximal capacity to dissipate body heat and therefore avoid the detrimental consequences of hyperthermia – the heat dissipation limit (HDL) theory. We contend that the HDL is a major constraint operating on the expenditure side of the energy balance equation, and that processes that generate heat compete and trade-off within a total boundary defined by heat dissipation capacity, rather than competing for limited energy supply.

    3. The HDL theory predicts that daily energy expenditure should scale in relation to body mass (Mb) with an exponent of about 0.63. This contrasts the prediction of the MTE of an exponent of 0.75.

    4. We compiled empirical data on field metabolic rate (FMR) measured by the doubly-labelled water method, and found that they scale to Mb with exponents of 0.647 in mammals and 0.658 in birds, not significantly different from the HDL prediction (P > 0.05) but lower than predicted by the MTE(P < 0.001). The same statistical result was obtained using phylogenetically independent contrasts analysis. Quantitative predictions of the model matched the empirical data for both mammals and birds. There was no indication of curvature in the relationship between Loge FMR and LogeMb.

    5. Together, these data provide strong support for the HDL theory and allow us to reject the MTE, at least when applied to endothermic animals.

    6. The HDL theory provides a novel conceptual framework that demands a reframing of our views of the interplay between energy and the environment in endothermic animals, and provides many new interpretations of ecological and evolutionary phenomena.

    Peer recognition: The paper was recommended at Faculty 1000 biology by Hans Otto Portner on May 26th 2010 as 'exceptional'. It was also recommended by Michael Symonds on the 10th June 2010 graded as a 'recommended read'.

    Highly cited: By January 2012 this paper was the 2nd highest cited paper from the 143 articles published by JAE during 2010.

    Independent follow-up studies

    A) The potential of the heat dissipation limitation theory to explain several phenomena in birds was explored by Gremillet et al (2012)Functional ecology 26: 1001-1006.

    Heat dissipation limit theory and the evolution of avian functional traits in a warming world

    Abstract: It is generally assumed that animal energy expenditure is limited by energy acquisition. In a series of publications, Speakman, Krol and colleagues argue that the capacity to dissipate metabolic heat may also limit maximum rates of energy expenditure in endotherms (heat dissipation limit theory HDL theory). The implications of the HDL theory for the evolution of avian functional traits are substantial and open fascinating research perspectives. Notably, the HDL theory leads us to (i) link elevated bird body temperatures with their capacity to achieve higher rates of heat loss and of energy expenditure, (ii) reconsider the evolution of avian plumage patterns and speculate upon the capacity of white birds to achieve higher field metabolic rates than darker relatives, (iii) hypothesize that the avian brood patch also functions as a thermal window allowing birds to shed excess heat and (iv) revise our current view of the adaptive significance of limited plumage thermal insulation in great cormorants. Such features have important implications for the capacity of birds to cope with global warming and for the design of mechanistic models of animal energetics aiming at predicting their responses to changing environmental conditions.

    B) In their comprehensive review of the factors driving metabolic allometry in birds and mammals Dan Reuman and colleagues came to the conclusion that among the available models the heat dissipation limitation theory provided the most convincing explanation of the observed patterns of variation in daily energy expenditure.

    The relationship between body mass and field metabolic rate among individual birds and mammals

    Hudson et al (2013) Journal of Animal Ecology 82: 1009-1020.


    1. The power-law dependence of metabolic rate on body mass has major implications at every level of ecological organization. However, the overwhelming majority of studies examining this relationship have used basal or resting metabolic rates, and/or have used data consisting of species-averaged masses and metabolic rates. Field metabolic rates are more ecologically relevant and are probably more directly subject to natural selection than basal rates. Individual rates might be more important than species-average rates in determining the outcome of ecological interactions, and hence selection.

    2. We here provide the first comprehensive database of published field metabolic rates and body masses of individual birds and mammals, containing measurements of 1498 animals of 133 species in 28 orders. We used linear mixed-effects models to answer questions about the body mass scaling of metabolic rate and its taxonomic universality/heterogeneity that have become classic areas of controversy. Our statistical approach allows mean scaling exponents and taxonomic heterogeneity in scaling to be analysed in a unified way while simultaneously accounting for nonindependence in the data due to shared evolutionary history of related species.

    3. The mean power-law scaling exponents of metabolic rate vs. body mass relationships were 0.71 [95% confidence intervals (CI) 0.625-0.795] for birds and 0.64 (95% CI 0.564-0.716) for mammals. However, these central tendencies obscured meaningful taxonomic heterogeneity in scaling exponents. The primary taxonomic level at which heterogeneity occurred was the order level. Substantial heterogeneity also occurred at the species level, a fact that cannot be revealed by species-averaged data sets used in prior work. Variability in scaling exponents at both order and species levels was comparable to or exceeded the differences 3/4-2/3 = 1/12 and 0.71-0.64.

    4. Results are interpreted in the light of a variety of existing theories. In particular, results are consistent with the heat dissipation theory of Speakman & Krol (2010) and provided some support for the metabolic levels boundary hypothesis of Glazier (2010).

    5. Our analysis provides the first comprehensive empirical analysis of the scaling relationship between field metabolic rate and body mass in individual birds and mammals. Our data set is a valuable contribution to those interested in theories of the allometry of metabolic rates.

    Supplementary information from our paper for download

    1. Model of heat loss as a function of size and ambient temperature in a resting mammal and bird (s1)
    2. Species data for field metabolic rate in mammals (s2)
    3. Species data for field metabolic rate in birds (s3)
  • {313}

    Shore, A., Karamitri, A., Kemp, P. SPEAKMAN, J.R. and Lomax, M.A (2010)

    Role of UCP1 enhancer methylation and chromatin remodelling in the control of UCP1 expression in adipose tissue

    Diabetologia 53:1164–1173

  • {312}

    SPEAKMAN, J.R. and Goran, M.I. (2010)

    Tissue-specificity and ethnic diversity in obesity related risk of cancer.

    Obesity 18: 1071-1078

  • {311}

    Hempenstall, S., Picchio, L., Mitchell, S.E., SPEAKMAN, J.R. and Selman, C. (2010)

    The impact of acute caloric restriction on the metabolic phenotype in male C57BL/6 and DBA/2 mice

    Mechanisms of Ageing and Development 131: 111-118

  • {310}

    SPEAKMAN, J.R. (2010)

    FTO effect on energy demand versus food intake

    Nature DOI: 10.1038/nature08807

    (brief communication arising: available on-line only)

  • {309}

    Duarte, L., Vaanholt, L., Sinclair, R., Gamo, Y., and SPEAKMAN, J.R. (2010)

    Limits to sustained energy intake XII: is the poor relationship between RMR and reproductive performance because RMR is not a repeatable trait?

    Journal of Experimental Biology 213: 278-287

  • {308}

    Larivée, M.L., Boutin, S., SPEAKMAN, J.R., McAdam, A.G. and Humphries, M.M. (2010)

    Associations between over-winter survival and resting metabolic rate in juvenile North American red squirrels (Tamiasciurus hudsonicus)

    Functional Ecology 24: 597-607

  • {307}

    Gerth, N., Redman, P., SPEAKMAN, J.R, Jackson, S. and Starck, J.M. (2010).

    Energy metabolism in Inuit sled dogs.

    Journal of Comparative physiology B 180: 577-589

  • {306}

    Banks, R., SPEAKMAN, J.R. and Selman, C. (2010)

    Vitamin E supplementation and mammalian lifespan.

    Molecular nutrition and Food research 54: 719-725.

  • {305}

    Welcker, J., Moe, B., Bech, C., Fyhn, M., Schultner, J., SPEAKMAN, J.R. and Gabrielsen, G. (2010)

    Evidence for an intrinsic energetic ceiling in free-ranging kittiwakes Rissa tridactyla.

    Journal of Animal Ecology 79: 205-213

  • {304}

    SPEAKMAN, J.R. and Westerterp, K.R. (2010)

    Reverse epidemiology, obesity and mortality in chronic kidney disease: modeling mortality expectations using energetics.

    Blood purification 29: 150-157

  • {303}

    Hinsley, S.A., Hill, R.A., Bellamy, P.E., Broughton, R.K., Harrison, N.M., Mackenzie, J.A., SPEAKMAN, J.R. and Ferns, P.N. (2009)

    Do Highly Modified Landscapes Favour Generalists at the Expense of Specialists? An Example using Woodland Birds.

    Landscape research 23: 615-626

  • {302}

    Reid, K.A., Margaritoulis, D and SPEAKMAN, J.R. (2009)

    Incubation temperature and energy expenditure during development in loggerhead sea turtle embryos

    Journal of Experimental Marine Biology and Ecology378: 62-68

  • {301}

    Sale, M., SPEAKMAN, J.R. and Arnould, J. (2009)

    Energy expenditure, water flux and activity budgets of Swamp Antechinuses in contrasting habitats.

    Journal of Mammalogy 90: 1238-1245

  • {300}

    Sheriff, M.J., SPEAKMAN, J.R., Kuchel, L., Boutin, S. and Humphries, M.M. (2009)

    The cold shoulder: free-ranging snowshoe hares maintain a low cost of living in cold climates.

    Canadian Journal of Zoology 87: 956-964.

  • {299}

    Lane, J.E., Boutin, S., SPEAKMAN, J.R., and Humphries, M.M. (2010)

    Energetic costs of male reproduction in a scramble competition mating system

    Journal of Animal Ecology 79:27-34

  • {298}

    Karamitri, A., Shore, A., Docherty, K., SPEAKMAN, J.R. and Lomax, M.A. (2009)

    Combinatorial transcription factor regulation of the cyclic response element on the PGC-1α promoter in white 3T3-L1 and brown HIB-1B preadipocytes.

    Journal of Biological Chemistry 284: 20738-20752

  • {297}

    Wu, S.H., Zhang, L., SPEAKMAN, J.R. and Wang, D.H. (2009)

    Limits to sustained energy intake XI: A test of the heat dissipation limit hypothesis in lactating Brandt’s voles (Lasiopodomys brandtii)

    Journal of Experimental Biology 212: 3455-3465.

  • {296}

    Fyfe, C.L., Stewart, J., Murison, S.D., Jackson, D.M., Rance, K., SPEAKMAN, J.R., Horgan, G.W. and Johnstone, A.M. (2010)

    Measuring food intake in free-living subjects - when to record and for how long?

    Public Health Nutrition 13: 172-180

  • {295}

    Djafarian, K., Jackson, D.M., Milne, E., Roger, P. and SPEAKMAN, J.R. (2010).

    Doubly-labeled water: Multi- and two-point methods in young children

    International Journal of Pediatric Obesity 5: 102-110.

  • {294}

    Welcker, J., Harding,A.M.A., Kitaysky, A., SPEAKMAN, J.R. and Gabrielsen, G.W. (2009)

    Daily energy expenditure increases in response to high food availability in an Arctic-breeding seabird, the little auk (Alle alle)

    Functional Ecology 23: 1081-1090

  • {293}

    Schmid, J. and SPEAKMAN, J.R. (2009)

    Torpor and energetic consequences in free ranging gray mouse lemurs (Microcebus murinus): a comparison of dry and wet forests.

    Naturwissenschaften 96: 609-20

  • {292}

    Jackson, D.M., Djafarian, K., Stewart, J., and SPEAKMAN, J.R. (2009)

    Increased TV viewing is associated with elevated body fatness but not lower energy expenditure.

    American Journal of Clinical Nutrition89: 1-6

  • {291}

    Williams, T.D., Vézina, F., & SPEAKMAN, J.R. (2009)

    Individually-variable energy management strategies during egg production are repeatable across broods.

    Journal of Experimental Biology212: 1101-1105

  • {290}

    Zub K., Szafrańska P. A., Konarzewski M., Redman P., SPEAKMAN, J.R. (2009)

    Tradeoffs between activity and thermoregulation in a small carnivore, the least weasel Mustela nivalis.

    Proceedings of the Royal Society B 276: 1921-1927

    This paper was featured on the cover of the issue

  • {289}

    SPEAKMAN, J.R. (2008)

    Thrifty genes for obesity and diabetes, an attractive but flawed idea and an alternative scenario: the ‘drifty gene' hypothesis.

    International Journal of Obesity 32: 1611-1617

    This paper was an invited perspective following the presidential lecture/debate presented at the Obesity Society annual meeting in New Orleans during November 2007. A sister paper by Andrew Prentice and colleagues which presents a view defending the Thrifty gene idea is published adjacent to this manuscript in the journal.

    see Prentice AM, Hennig BJ, Fulford AJ. Evolutionary origins of the obesity epidemic: natural selection of thrifty genes or genetic drift following predation release? Int J Obes (Lond). 32: 1606-1610

    See also independent review of this paper in

    Robson, S (2008) Some insights into the epidemiology of obesity. O&G magazine summer 2008 pp 37-38

    Also note internet reviews of this paper

    Carlson Chow Genetic basis of obesity

    This work was subject of a radio interview at WHBC Ithaca (2/2/2011) when I gave a presentation on it for the McCay lectureship at Cornell University in February 2011

    Drifty gene on wikipedia

  • {288}

    Kelm, D.H., Schaer, J., Ortmann, S., Wibbelt, G., SPEAKMAN, J.R., and Voigt, C.C. (2008)

    Efficiency of facultative frugivory in the nectar-feeding bat Glossophaga commissarisi: the quality of fruits as an alternative food source

    Journal of Comparative Physiology B 178: 985-996.

  • {287}

    Voigt, C.C., Kretzschmar, A.S., SPEAKMAN, J.R. and, Lehmann, G.U.C. (2008). Female bushcrickets fuel their metabolism with male nuptial gifts. Biology Letters 4: 476-478

    The paper was recommended at Faculty 1000 Biology by Mike Ritchie on July 7th 2008 graded as a "recommended read".

  • {286}

    Voigt, C.C., Baier, L., SPEAKMAN, J.R. and Siemers, B.M. (2008)

    Stable carbon isotopes in exhaled breath as tracers for dietary information in birds and mammals

    Journal of Experimental Biology 211: 2233-2238

  • {285}

    Westerterp, K.R. and SPEAKMAN, J.R. (2008)
    Physical activity energy expenditure has not declined since the 1980s and matches energy expenditure of wild mammals

    International Journal of Obesity 32:1256-63

    media coverage and impact word doc

    Brief summary Objective measurements of energy expenditure using doubly labelled water between 1985 and 2005 show no trend over time. The same is true for various measures of physical activity energy expenditure. Moreover the estimates do not differ from those in the third world or the levels predicted from wild animals. Contrary to popular belief objective measures of expenditure do not support the idea that physical activity energy expenditure has declined - at least back to the 1980s.

    Chinese summary: 从1985年到2005年间使用双标水法对能量耗散进行的客观测量并不过时。各种对身体活动能量耗散进行测量的方法也是同样的道理。此外,这些测量值和第三世界或者和野生动物预测的水平没有什么不一样。和流行的看法相反的是-对能量耗散客观测量的结果并不支持身体活动能量耗散下降的说法(至少80年代到现在)。


    Objective: Obesity results from protracted energy imbalance. Whether this comprises excessive energy intake, lowered physical activity or both, remains disputed.

    Design: Physical activity energy expenditure, evaluated in three different ways from daily energy expenditure (DEE) measured using doubly labelled water, was examined for trends over time. Data included subjects in Europe (Maastricht, the Netherlands) and North America extending back to the 1980s. These data were compared with measures from the third world, and measures made on wild terrestrial mammals.

    Results: Physical activity expenditure in Europe (residual of the regression of DEE on basal energy expenditure (BEE)) has slightly but significantly increased since the 1980s. There was no trend over time in physical activity level (PAL = DEE/BEE), or in the residual variance in DEE once mass, sex and age were accounted for. This latter index of physical activity expenditure also significantly increased over time in North America. DEE of individuals in Europe and North America was not significantly different from individuals measured in the third world. In wild terrestrial mammals, DEE mostly depended on body mass and ambient temperature. Predicted DEE for a 78 kg mammal living at 20 oC was 9.2 MJ per day (95% CI: 7.9–12.9 MJ per day), not significantly different from the measured DEE of modern humans (around 10.2–12.6 MJ per day).

    Conclusion: As physical activity expenditure has not declined over the same period that obesity rates have increased dramatically, and daily energy expenditure of modern man is in line with energy expenditure in wild mammals, it is unlikely that decreased expenditure has fuelled the obesity epidemic.

    Peer recognition: The paper was recommended at Faculty 1000 medicine by Mikael Fogelholm on July 7th 2008 graded 3.0 as a recommended read. It was also recommended at faculty 1000 Biology by Hannah Carey on 9th January 2009, graded 3.0 as a recommended read.

    Highly cited: By January 2012 this paper was the 16th highest cited paper from 1284 articles published by the IJO during 2008 (top 2%).

  • {284}

    SPEAKMAN, J.R., Rance, K.A. and Johnstone, A.M. (2008)
    Polymorphisms of the FTO gene are associated with variation in energy intake but not energy exprenditure.

    Obesity 16:1961-5

    media coverage and impact word doc

    Brief summary First study to show polymoprhisms in the SNP rs 9939609 near to the FTO gene are linked to food intake (and protein intake) variation and not energy expenditure.

    Chinese summary 对FTO基因附件的SNP rs 9939609 多态性和食物摄入(蛋白质摄入)变化相关和不是能量的支出的第一个研究。


    The FTO gene has significant polymorphic variation associated with obesity, but its function is unknown. We screened a population of 150 whites (103F/47M) resident in NE Scotland, United Kingdom, for variants of the FTO gene and linked these to phenotypic variation in their energy expenditure (basal metabolic rate (BMR) and maximal oxygen consumption VO2max) and energy intake. There was no significant association between the FTO genotype and BMR or VO2max. The FTO genotype was significantly associated (P = 0.024) with variation in energy intake, with average daily intake being 9.0 MJ for the wild-type TT genotype and 10.2 and 9.5 MJ for the “at risk” AT and AA genotypes, respectively. Adjusting intake for BMR did not remove the significance (P = 0.043). FTO genotype probably affects obesity via effects on food intake rather than energy expenditure.

    Highly cited: By January 2012 this paper was the 4th highest cited paper from 1424 articles published by Obesity during 2008 (top 0.5%). It was also the 21st highest cited paper ever published by the journal from a total of 5921 articles (top 0.5%).

  • {283}

    Selman, C., McLaren, J.S., Collins, A.R., Duthie, G.G. and SPEAKMAN, J.R. (2008)

    The impact of experimentally elevated energy expenditure on oxidative stress and lifespan in the short-tailed field vole Microtus agrestis.

    Proceedings of the Royal Society of London B 275:1907-16.

    see also independent review of this paper in

    Darveau C Outside JEB section Journal of Experimental biology 211: ppV

  • {282}

    Voigt, C.C., Rex, K., Michener, R.H. and SPEAKMAN, J.R. (2008)

    Nutrient routing in omnivorous animals tracked by stable carbon isotopes in tissue and exhaled breath

    Oecologia 157: 31-40.

  • {281}

    Furness, L. and SPEAKMAN, J.R. (2008)

    Energetics and longevity in birds.

    AGE 30: 75-87

  • {280}

    Hinsley, S.A., Hill, R.A., Bellamy, P.E., Harrison, N.M., SPEAKMAN, J.R., Wilson, A.K. and Ferns, P.N. (2008)

    Effects of structural and functional habitat gaps on breeding woodland birds: working harder for less

    Landscape Ecology 23: 615-626

  • {279}

    Simončič, M., Horvat, S., Stevenson, P.L., Bünger, L., Holmes, M.C., Kenyon,C.J., SPEAKMAN, J.R. and Morton, N.M. (2008).

    Divergent physical activity and novel alternative responses to high fat feeding in polygenic fat and lean mice.

    Behaviour genetics 38: 292-300

  • {278}

    Tidhar, W.L. and SPEAKMAN, J.R. (2007)

    An evaluation of four non-destructive methods for predicting body composition of a small rodent.

    International Journal of Body Composition Research 5: 137-145

  • {277}

    Sparling, C.E., Thompson, D., Fedak, M.A., Gallon, S.M. and SPEAKMAN, J.R. (2008)

    Estimating the field metabolic rate of pinnipeds: doubly-labelled water gets the seal of approval.

    Functional Ecology 22: 245-254.

    This paper was awarded the Haldane prize for the best paper published in Functional Ecology during 2008 that was first authored by a young scientist.

  • {276}

    SPEAKMAN, J.R., Hambly, C., Mitchell, S.E. & Krol, E. (2008)
    The contribution of animal models to the study of obesity
    Laboratory animals 42: 413-432

    This paper is an extended version of an invited paper that was part of the Government foresight review "Tackling obesities: Future choices" compiled by Sir David King. The entire report of this project (12Mb) can be downloaded here

    individual invited contributions were published in a supplment to Obesity Reviews. Our contribution was

    SPEAKMAN, J.R., Hambly, C., Mitchell, S.E. and Krol, E. (2008) Animal models of obesity. Obesity reviews 8: supplement 1 55-61.

  • {275}

    Haggerty, C., Hoggard,N., Brown,D.S., Clapham, J.C. and SPEAKMAN, J.R. (2008)

    Intra-specific variation in resting metabolic rate in MF1 mice is not associated with membrane lipid desaturation in the liver

    Mechanisms of Ageing and Development 129: 129-137

  • {274}

    Adams, A.C., Wynick, D., Clapham, J.C. and SPEAKMAN, J.R. (2008)

    Feeding behavior in galanin knockout mice supports a role of galanin in fat intake and preference

    Journal of Neuroendocrinology 20: 199-206

  • {273}

    Krol, E., Murphy, M. and SPEAKMAN, J.R. (2007)

    Limits to sustained energy intake X: Effects of fur removal on reproductive performance in laboratory mice

    Journal of Experimental Biology 207: 4233-4243

    This paper was a subject of an ‘inside JEB' article in the same issue

    Phillips, K. (2007) Limits on lactation. JEB iii

    and an article in New Scientist (24th Nov 07)

    Shaving may boost milk production in mums

    It was a featured interview on the Canadian national radio program ‘Quirks and Quarks' Nov 07

    It was also featured on several online science news sites in November 2007

    • New scientist ( - ‘Naked' mums produce plumper babies
    • Thaindian news ( - Naked mums produce chubbier offspring
    • Digg ( - Naked mums produce chubbier babies
    • Yahoo news ( - Naked mums raise plumper babies
    • Webindia ( - Naked mums raise chubbier offsprings
    • The cheers mag news ( - Naked mums raise chubbier offspring
    • Daily India ( - Naked mums produce chubbier offspring
    • Softpedia ( - Naked mother for stronger toddlers
    • Zigg ( - 'Naked' mums produce babies plumpest
    • e-biology news ( - Naked mums raise plumper offsprings
    • Top news ( - Naked mums raise chubbier offsprings
    • Mylder ( - Naked mums produce plumper babies
    • Word press ( - naked mums produce more milk
    • Inform ( - shaving may boost milk production in mums
    • Newsbrief ( - Naked mums produce plumper babies
    • Scirus ( - naked mums produce plumper babies
  • {272}

    Selman, C., Lingard, S, Choudhury, A.I., Batterham, R.L., Claret, M., Clements, M., Ramadani, F., Okkenhaug, K., Schuster, K., Blanc, E., Piper, M.D., Al-Qassab, H., SPEAKMAN, J.R., Carmignac, D., Robinson, I.C.A., Thornton, J.M., Gems, D., Partridge, L. and Withers, D.J. (2008)

    Evidence for lifespan extension and delayed age related biomarkers in insulin receptor substrate 1 null mice. FASEB J. 22: 807-818

    This paper was reported in the press in Oct 2007 when the e-pub was released

    The Guardian - Lifespan gene in mice could help restrict old age diseases
    Hindustan times - Scientists identify lifespan gene
    India Daily News - Gene that regulates lifespan found

    And on several science web sites - A longer-living, healthier mouse that could hold clues to human ageing
    BS online news - Scientists discover gene that regulates lifespan

    Highly cited: by January 2012 this was the 3rd highest cited paper from a total of 442 published by FASEB J in 2008.

  • {271}

    Valle, A., Hoggard, N., Adams, A.C., Roca , P., and SPEAKMAN, J.R. (2008)

    Chronic central administration of apelin-13 over ten days increases food intake, body weight, locomotor activity and body temperature in C57BL/6 mice.

    Journal of Neuroendocrinology 20: 79-84.

  • {270}

    Wallach, A.D., Inbar, M., Scantlebury, M., SPEAKMAN, J.R. and Shanas, U. (2007)

    Water Requirements as a Deer Reintroduction Bottleneck at the Southern Edge of its Range

    Canadian Journal of Zoology 85: 1182-1192

  • {269}

    Selman, C., McLaren, J.S., Mayer, C., Duncan, J.S., Collins, A.R., Duthie, G.G., Redman, P., and SPEAKMAN, J.R. (2008)

    Lifelong α-tocopherol supplementation increases median lifespan of C57BL/6 mice in the cold but has only minor effects on oxidative damage.

    Rejuvenation Research 11: 83-95.

  • {268}

    Harris, M.P., Newall, M., Daunt, F., SPEAKMAN, J.R. and Wanlass, S. (2008)

    Snake pipefish Entelurus aequoreus are poor food for seabirds

    Ibis 150: 413-415

  • {267}

    Hoggard, N., Bashir, S., Cruickshank, M., Miller J.D.B. and SPEAKMAN, J.R. (2007)

    Expression of Neuromedin B in adipose tissue and its regulation by changes in energy balance.

    J. Mol. Neuroendocrinology 39: 199-210.

  • {266}

    Vaanholt, L.M., SPEAKMAN, J.R., Garland T., Lobley, G.E., and Visser, G.H. (2008)

    Protein synthesis and antioxidant capacity in ageing mice: effects of long-term voluntary exercise.

    Physiological and Biochemical Zoology 81: 148-157.

    This paper was listed on the PBZ web site in November 2008 as the 5th most downloaded paper from their site.

  • {265}

    Voigt, C.C., Grasse, P., Rex, K., Hetz, S.K. and SPEAKMAN, J.R.(2007)
    Bat breath reveals metabolic substrate use in free-ranging vampires.
    Journal of Comparative Physiology B 178: 9-16.

    This paper was widely reported on online science news sites in August 2007

    • - Vampire bats attacking cattle as rain forest falls
    • - Bat breath reveals identity of a vampires last victim
    • - Bat breath reveals identity of vampires last victims
    • - Costa Rican vampire bats thrive on cattle blood
    • - Costa Rican vampire bats thrive on cattle blood
    • - Bite of the vampire: bat breath gives clue to shifting Latin American environment
    • - Clearing rainforest for cattle pastures drives surge in vampires
    • - Bite of the vampire
    • - Thriving on cattle blood vampire bats proliferate
    • - Fresh cattle blood is booming the numbers of vampires
    • - Ecology of vampire bats
    • - Ecological changes forcing vampire bats to attack cattle
    • - Bat breath reveals identity of a vampires last victim
    • - Bite of the vampire
    • - Thriving on cattle blood vampire bats proliferate
    • - Vampires overtake costa rica
    • - Vampires thriving on cattle blood
    • - Vampir-Atem verr?t die Identit?t des letzten Opfers
    • - Vampir-Atem verr?t die Identit?t des letzten Opfers
    • - Vampir-Atem verr?t die Identit?t des letzten Opfers
    • - bat breath reveals identity of a vampires last victim
    • - Tell tale bat breath
    • - Vampires switch to cattle
    • - Vampire bats and stable isotopes
    • - Ecological changes forcing bats to attack cattle
  • {264}

    Hambly, C., Mercer, J.G. and SPEAKMAN, J.R. (2007)

    Hunger does not diminish over time in mice under protracted caloric restriction.

    Rejuvenation Research 10: 533-540.

  • {263}

    Hambly, C., Simpson, C.A., McIntosh, S., Duncan, J.,Dalgleish G.D., and SPEAKMAN, J.R. (2007)

    Calorie-restricted mice that binge show less ability to compensate for reduced energy intake

    Physiology and Behaviour 95: 982-992.

  • {262}

    Voigt, C.C. and SPEAKMAN, J.R. (2007)
    A mammalian nectar specialist fuels its metabolically expensive nocturnal life directly and almost exclusively with exogenous carbohydrates
    Functional Ecology

    See review of this paper at, 15th August 2007

    See also articles on this paper in

    This paper was widely reported on online science web sites in August 2007

    • - Sugar rushes keep bats airborne
    • - Why nectar feeding bats need a power drink to fly
    • - Bats can gorge on sweets guilt free
    • - Nectar feeding bats are powered by pure sugar-
    • - Bats burn sugar better than other mammals
    • - Bats burn sugar faster than any other mammal on earth
    • - Fastest sugar burning mammals on earth
    • - Bats burn sugar better than other mammals
    • - Bats burn sugar at a phenomenal rate
  • {261}

    Bennett, K.A., SPEAKMAN, J.R., Moss, S., Pomeroy, P. and Fedak, M.A. (2007)

    Effects of mass and body composition on fasting fuel utilization in grey seal pups (Halichoerus grypus; Fabricius): an experimental study using supplementary feeding.
    Journal of Experimental Biology 210: 3043-53

  • {260}

    SPEAKMAN, J.R. (2007)

    A novel non-adaptive scenario explaining the genetic pre-disposition to obesity: the ‘predation release’ hypothesis.

    CELL metabolism 6: 5-11

    This paper was the most downloaded paper at Cell metabolismduring July 2007 and the second most downloaded paper during August 2007.

    The paper was recommended at Faculty 1000 Biology by Andries Karlsbeek on August 16th 2007 graded as a "Must read". See

  • {259}

    Rance, K.A., Johnstone, A.M., Murison. S., Duncan, J.S., Wood, S.G. and SPEAKMAN, J.R. (2007)

    Plasma leptin levels re related to body composition, sex, insulin levels and the A55V polymorphism of the UCP2 gene.

    International Journal of Obesity

  • {258}

    Tidhar, W.L., Bonier, F., and SPEAKMAN, J.R. (2007)

    Sex- and concentration-dependent effects of predator feces on seasonal regulation of body mass in the bank vole Clethrionomys glareolus

    Hormones and Behaviour 52: 436-444

  • {257}

    Scantlebury, M., Waterman, J.M., Hillegrass, M., SPEAKMAN, J.R. & Bennett, N.C. (2007).

    Energetic costs of parasitism in the Cape ground squirrel Xerus inauris

    Proceedings of the Royal Society of London B 274: 2169-2177

  • {256}

    Rance, K.A., Hambly, C., Dalgleish, D., Fustin, J.M., Bünger, L., and SPEAKMAN, J.R. (2007)

    Quantitative Trait Loci for total and regional adiposity in mouse lines divergently selected for food intake

    Obesity 15: 2994-3004.

  • {255}

    Claret, M., Smith, M.A., Batterham, R.L., Selman, C., Choudhury, A.I., Fryer, L.G.D., Clements, M., Al-Qassab, H., Heffron, H., Xu, A.W., SPEAKMAN, J.R., Barsh, G.S., Viollet, B., Vaulont, S., Ashford, M.L.J., Carling, D. and Withers, D.J. (2007)

    AMPK is essential for energy homeostasis regulation and glucosesensing by POMC and AgRP Neurons.

    Journal of Clinical Investigation 117: 2325-2336

  • {254}

    SPEAKMAN, J.R., Djafarian, K., Stewart, J. and Jackson, D.M. (2007)

    Assortative mating for obesity

    American Journal of Clinical Nutrition 86: 316-23

    Brief summary: study of 42 couples in scotland shows assortative mating for total and regional aspects of adiposity and body composition measured using DXA, confirming previous studies based only on BMI. Theoretical model shows how increase in assortative mating may have contributed to the obesity epidemic.


    Background: Assortative mating is the nonrandom mating of individuals with respect to phenotype and cultural factors. Previous studies of assortative mating for obesity have indicated that it may have contributed to the obesity epidemic. However, those studies all used body mass index or skinfold thicknesses to measure obesity and did not always account for potential confounding factors.

    Chinese summary: .通过对42对夫妇苏格兰用DEXA进行整体和部分体脂和身体成分的测量的选择性择偶的研究证实了之前仅仅基于BMI的研究结果。理论模型表明了选择性择偶的增加是如何影响肥胖的趋势的。

    Objective: We aimed to assess the level of assortative mating for obesity by using dual-energy X-ray absorptiometry to characterize body composition.

    Design: This was a cross-sectional study of 42 couples.

    Results: Raw spousal correlations showed assortative mating for age, weight, body mass index, lean mass, and fat mass. Removing the effect of age on fat mass strengthened the spousal correlation (r = 0.405). Social homogamy did not appear to be important, because in this sample there was no significant effect of area of origin on age-corrected fat and lean tissue masses for either sex. Regional body-composition analysis showed that subjects with disproportionately large arms (both fat and lean) assortatively mated with partners with the same trait. However, both men and women with high lean tissue in their arms assortatively mated with partners that had a disproportionately low fat content in their legs.

    Conclusions: These data confirm that assortative mating for obesity exists when dual-energy X-ray absorptiometry is used to evaluate adiposity. We hypothesize that assortative mating may have contributed to the obesity epidemic because the time course of obesity development has shifted progressively earlier, allowing singles in their late teens and early twenties to more easily distinguish partners with obese and lean phenotypes.

  • {253}

    Aquarone, M., Born, E.W. and SPEAKMAN, J.R. (2006)

    Field metabolic rates of Walrus (Odobenus rosmarus) measured by the doubly-labelled water method.

    Aquatic mammals 32: 363-369.

  • {252}

    Stocker, C.J., Wargent, E., O’Dowd, J., Cornick, C., SPEAKMAN, J.R., Arch, J.R., and Cawthorne, M.A. (2007)

    Prevention of diet-induced obesity and impaired glucose tolerance in rats following administration of leptin to their mothers.

    Am. J. Physiol. 282: R1810-1818

  • {251}

    Vezina, F., SPEAKMAN, J.R., and Williams, T.D. (2006)

    Individually variable energy management strategies in relation to the costs of egg production

    Ecology 87: 2447-2458

  • {250}

    Jackson, D.M., Pace, L. and SPEAKMAN, J.R. (2007)

    The measurement of RMR in pre-school children.

    Obesity 15: 1930-1932

  • {249}

    Tolkamp, B., Yearsley, J., Gordon, I., Illius, A., SPEAKMAN, J.R. and Kyriazakis, I. (2007

    Predicting the effects of body fatness on food intake and performance of sheep.

    British Journal of Nutrition.21: 1-10

  • {248}

    Visockiene, Z., Johnstone, A.M., SPEAKMAN, J.R. and Broom, J.I. (2007)

    Body composition measurement in obese patients with and without type 2 diabetes:comparison of methods

    Acta medica Lithuanica14: 301-308

  • {247}

    Król, E., Tups, A., Archer, Z.A., Ross, A.W., Moar, K.M., Bell, L.M., Duncan, J.S., Mayer, C., Morgan, P.J., Mercer, J.G. and SPEAKMAN, J.R. (2007)

    Altered Expression of SOCS3 in the Hypothalamic Arcuate Nucleus during Seasonal Body Mass Changes in the Field Vole, Microtus agrestis.

    Journal of Neuroendocrinology 19: 83-94

  • {246}

    SPEAKMAN, J.R. and Hambly, C. (2007)

    Starving for life – what animal studies can, and cannot, tell us about the use of caloric restriction to prolong human lifespan.

    Journal of Nutrition 137:1078-1086

  • {245}

    Johnston, S.J., Souter, D.M., Erwin, S.S., Tolkamp, B.J., Yearsley, J.M., Gordon, I.J., Illius, A., Kyriazakis, I. and SPEAKMAN, J.R. (2007)

    Associations between basal metabolic rate and reproductive performance in C57BL/6J mice

    Journal of Experimental Biology 210: 65-74

  • {244}

    Krol, E. and SPEAKMAN, J.R. (2007)

    Regulation of body mass and adiposity in the field vole, Microtus agrestis: a model of leptin resistance

    Journal of Endocrinology 192: 271-278

  • {243}

    Kokurewicz, T. and SPEAKMAN, J.R. (2007)

    Age related effects on energy metabolism during torpor in Daubenton’s bats: effects on fat accumulation prior to hibernation.

    Acta Chiropterologica 8: 509-521

  • {242}

    Rance, K.A., Johnstone, A.M., Murison, S., Duncan, J.S., Wood, S.G.,and SPEAKMAN, J.R. (2007)

    Circulating leptin levels are related to adiposity, age, sex and the A55V polymorphism of the UCP2 gene.

    International Journal of Obesity 31: 1311-1318

  • {241}

    Johnston, S.L., Erwin, S.S., Souter, D.M., Tolkamp, B.J., Gordon, I.J., Illius, A.W., Kyriazakis, I. and SPEAKMAN, J.R (2007)

    Intake compensates for resting metabolic rate variation in female C57BL/6J mice fed high-fat diets.

    Obesity 15: 600-606

  • {240}

    SPEAKMAN, J.R. (2007)

    The energy cost of reproduction in small rodents.

    Acta theriologica sinica 27: 1-13

  • {239}

    Selman, C., McLaren, J.S., Meyer, C., Duncan, J.S., Redman, P., Collins, A.R., Duthie, G.G. and SPEAKMAN, J.R. (2006)

    Life-long vitamin C supplementation in combination with cold exposure does not affect oxidative damage or lifespan in mice, but decreases expression of antioxidant protection genes

    Mechanisms of Ageing and Development 127: 897-904

  • {238}

    Visockiene, Z., Broom, J.I., SPEAKMAN, J.R. and Johnstone, A.M. (2006)

    Dietary intake and self-reporting in relation to eating behaviour in obese and type 2 diabetes patients

    Baltic endocrinology 2:24-30

  • {237}

    SPEAKMAN, J.R. (2008)

    The physiological cost of reproduction in small mammals.

    Philosophical transactions of the Royal Society 363: 375-398

  • {236}

    Sparling, C.E., SPEAKMAN, J.R. and Fedak, MA. (2006)

    Seasonal variation in the metabolic rate and body composition of female grey seals: fat conservation prior to high-cost reproduction in a capital breeder?

    Journal of Comparative Physiology B 176: 505-512

  • {235}

    Johnstone, A.M., Rance, K.A., Murison, S.D., Duncan, J.S. and SPEAKMAN, J.R. (2006)

    Taking additional anthropometric measures may improve the predictability of Basal Metabolic Rate in adult subjects

    European Journal of Clinical Nutrition 60: 1437-1444.

  • {234}

    SPEAKMAN, J.R. (2006)

    Thrifty genes for obesity and the metabolic syndrome – time to call off the search?

    Diabetes and vascular disease research 3: 7-11

  • {233}

    Arch, J.R.S., Hislop, D., Wang, S.J.Y., and SPEAKMAN, J.R. (2006)

    Some mathematical and technical issues in the measurement and interpretation of open-circuit indirect calorimetry in small animals

    International Journal of Obesity 30: 1322-1331

  • {232}

    Scantlebury, M., SPEAKMAN, J.R., Oosthuizen, M.K., Roper, T., & Bennett, N.C. (2006)

    Energetics reveals physiologically distinct castes in a eusocial mammal.

    Nature.440: 795-797

  • {231}

    Southwood, A.L., Reina, R.D., Jones, V.S., SPEAKMAN, J.R., and Jones, D.R. (2006).

    Seasonal metabolism of juvenile green turtles (Chelonia mydas) at Heron Island, Australia.

    Canadian Journal of Zoology

  • {230}

    Humphries M.M., Boutin S., Thomas D.W., Ryan J.D., Selman C., McAdam A.G., Berteaux D., SPEAKMAN, J.R. (2005)

    Expenditure freeze: the metabolic response of small mammals to cold environments

    Ecology letters 8: 1326-1333

  • {229}

    Johnston, S., Grune, T., Bell, L., Murray, S., Souter, D., Erwin, S., Yearsley, J., Gordon, I., Illius, A., Kyriazakis, I., SPEAKMAN, J.R. (2006)

    Having it all - historical energy intakes do not generate the anticipated trade-offs in fecundity.

    Proceedings of the Royal Society of London B 273: 1369-1374

  • {228}

    Judge S, Jang YM, Smith A, Selman C, Phillips T, SPEAKMAN, J.R., Hagen T, Leeuwenburgh C. (2005)

    Exercise by lifelong voluntary wheel running reduces subsarcolemmal and interfibrillar mitochondrial hydrogen peroxide production in the heart.

    American Journal of Physiology 289: R1564-1572.

  • {227}

    Yearsley, JM., Villalba, J., Kyriazakis, I., SPEAKMAN, J.R., Tolkamp, B.J., Illius, J.W., Gordon, I.J. and Duncan, A. (2005).

    A theory of associating food types with their post-ingestive consequences.

    American Naturalist 167: 705-716

  • {226}

    Król, E., Duncan, J.S., Redman, P., Morgan, P.J., Mercer, J.G. and SPEAKMAN, J.R. (2005)

    Photoperiod regulates leptin sensitivity in field voles, Microtus agrestis

    Journal of Comparative Physiology B 176: 153-163

  • {225}

    Mathias, ML., Nunes, A.C., Marques, C., Auffray, J.C., Britton Davidian, J., Catalan, J., Ganem, G., Grunduz, I., Ramathinho, M.G., Searle, J.B. and SPEAKMAN, J.R. (2006)

    Effects of climate on oxygen consumption and energy intake of chromosomally divergentpopulations of the house mouse (Mus musculus domesticus) from the island of Madeira (North Atlantic, Portugal).

    Functional Ecology 20: 330-339

  • {224}

    Scantlebury, M., Bennett, N.C., SPEAKMAN, J.R., Pillay, N. and Schradin, C. (2006) The energetics of huddling in group-living African four-striped field mice Rhabdomys pumilio

    Functional Ecology 20: 166-173

  • {223}

    Scantlebury, M., SPEAKMAN, J.R., and Bennett, N. (2006)

    The costs of sexual dimorphism in mole rats are morphological not behavioural.

    Proceedings of the Royal Society Series B: Biologial Sciences 273: 57-63.

  • {222}

    Johnston, S.L., Peacock, W.L., Bell, L.M., Lonchampt, M. and SPEAKMAN, J.R. (2005)

    PIXImus DXA with Different Software Need Individual Calibration to Accurately Predict Fat Mass

    Obesity Research 13: 1558-1565

  • {221}

    Johnstone, A.M., Murison, S.D., Duncan, J.S., Rance, K.A. and SPEAKMAN, J.R (2005)

    Factors influencing variation in basal metabolic rate include fat-free mass, fat mass, age, and circulating thyroxine but not sex, circulating leptin, or triiodothyronine.

    American Journal of Clinical Nutrition.82: 941– 8.

  • {220}

    Hambly, C.and SPEAKMAN, J.R. (2005)

    Contribution of different mechanisms to compensation for energy restriction in the mouse.

    Obesity Research 13: 1548-1557

  • {219}

    SPEAKMAN, J.R. and Krol, E. (2005) Limits to sustained energy intake IX: a review of hypotheses.

    Journal of Comparative Physiology B 175: 375-394

  • {218}

    SPEAKMAN, J.R., Walker, H., Walker, L., and Jackson, D.M. (2005)

    Associations between BMI, social strata and the estimated energy content of foods.

    International Journal of Obesity29: 1281-1288

  • {217}

    Selman, C., Phillips, T., Staib, J.L., Duncan, J.S., Leeuwenburgh, C., and SPEAKMAN, J.R. (2005)

    Energy expenditure of calorically restricted rats is higher than predicted from their altered body composition

    Mechanisms of Ageing and Development. 126: 783-793

    See also independent review of this paper in ?Outside JEB? Journal of Experimental Biology June 2005 By T. Valencak.

  • {216}

    Wanlass, S., Harris, M.P., Redman, P. and SPEAKMAN, J.R. (2005)

    Low energy values of fish as a probable cause of a major seabird breeding failure in the North Sea

    Marine Ecology progress series.294: 1-8

    This paper was featured on the front cover

  • {215}

    SPEAKMAN, J.R. (2005)

    Correlations between physiology and lifespan – two widely ignored problems with comparative studies.

    Aging Cell 4: 167-175

  • {214}

    Rance,K.A., Fustin, J.M., Dalgleish, G., Hambly, C., Bünger,L. and SPEAKMAN, J.R. (2005)

    A Paternally Imprinted QTL for Mature Body Mass on Mouse Chromosome 8

    Mammalian genome 16: 567-577

  • {213}

    Simon, A., Thomas, D.W., SPEAKMAN, J.R., Blondel, J., Perret, P. and Lambrechts, M. (2005) Impact of ectoparasitic blowfly larvae (Protocalliphora spp) on the behaviour and energetics of nestling blue tits (Parus caeruleus).

    Journal of Field Ornithology 76: 402-410

  • {212}

    Scantlebury, M., M.K. Oosthuizen SPEAKMAN, J.R., C.R. Jackson and Bennett, N.C.(2005)

    Seasonal energetics of the Hottentot golden mole (Ambysomus hottentottus longiceps) at High altitude (1500m)

    Physiology and behaviour 84: 739-745

  • {211}

    Choudhury, A.I., Heffron, H., Smith, M.A., Al-Qassab, H., Xu, A.W., Selman, C., Simmgen, M., Clements, M., Claret, M., MacColl, G., Hisadome, K., Diakonov, I., Moosajee, V., Bell, J.D., SPEAKMAN, J.R., Batterham, R.L., Barsh, G.S., Ashford, M.L.J. and Withers, D.J. (2005)

    The role of insulin receptor substrate 2 in hypothalamic and beta cell function

    Journal of Clinical Investigation 115: 940-950

  • {210}

    SPEAKMAN, J.R. (2005) The role of technology in the past and future development of the doubly-labelled water method. Isotopes in Environmental and health studies.41: 335-343.

  • {209}

    SPEAKMAN, J.R. (2005)

    Body size, energy metabolism and lifespan

    Journal of Experimental Biology 208: 1717-1730

    Highly cited: By November 2007 this paper was the 5th highest cited paper that had been published during 2005 by JEB, from a total of 599 articles published that year. By January 2012 it was the 4th highest cited article published by JEB in 2005 (top 1%). It was also the 12th highest cited paper from a total of 10000 papers published by JEB between 2002 and 2012

  • {208}

    Hambly, C., Adams, A.C., Fustin, J.M., Rance, K.A., Bünger, L. and SPEAKMAN, J.R. (2005).

    Mice with low metabolic rates are not susceptible to weight gain when fed a high fat diet.

    Obesity Research 13: 556-566

  • {207}

    Król, E., Redman, P., Thomson, P.J., Williams, R., Mayer, C., Mercer, J.G., and SPEAKMAN, J.R. (2005)

    Effect of photoperiod on body mass, food intake and body composition in the field vole, Microtus agrestis

    Journal of Experimental Biology 208: 571-584

  • {206}

    Hill, R.C., Lewis, D.D., Randell, S.J., Scott, K,C., Omori, M. Sundstrom, D.A., Jones, G.L., SPEAKMAN, J.R., and Butterwick, R.F. (2005)

    Mildly restricting food intake increases the speed of racing greyhounds

    American Journal of veterinary research 66: 1065-1070

  • {205}

    Ward, S., Moller, U., Rayner, J.M.V., Jackson,D.M., Nachtigall, W., and SPEAKMAN, J.R. (2004)

    Metabolic power of European Starlings (Sturnus vulgaris) during flight in a wind tunnel measured by heat transfer modeling, doubly-labelled water, mask respirometry and aerodynamic modeling,

    Journal of Experimental Biology 207: 4291-4298

  • {204}

    Yearsley, J.M., Kyriazakis, I., Gordon, I.J., Illius, A.W., Johnston, S.L., SPEAKMAN, J.R. and Tolkamp, B.J. (2005)

    A life history model of somatic damage associated with resource acquisition: damage protection or prevention?

    Journal of Theoretical Biology 235: 305-317

  • {203}

    Hambly, C., Harper, E.J., and SPEAKMAN, J.R. (2004)

    The energetic cost of variations in wing span and wing asymmetry in the zebra finch (Taeniopygia guttata).

    Journal of Experimental Biology207: 3977-3984
  • {202}

    Hambly, C., Harper, E.J., and SPEAKMAN, J.R. (2004)

    The energy cost of loaded flight is substantially lower than expected due to alterations in flight kinematics.

    Journal of Experimental Biology 207: 3969-3976

  • {201}

    Hambly, C., Pinshow, B., Wiersma, P., Verhulst, S., Piertney, S.B., Harper, E.J. & SPEAKMAN, J.R. (2004)

    Comparison of the cost of short flights in a nectarivorous and non-nectarivorous bird.

    Journal of Experimental Biology 207: 3959-3968
  • {200}

    Hoggard, N., Rayner, D.V., Johnston, S.L. and SPEAKMAN, J.R.(2004)

    Peripherally administered [Nle4, D-Phe7]-aMSH increases resting metabolic rate, while peripheral AgRP has no effect, in wild type C57BL/6 and ob/obmice.

    Journal of Molecular Endocrinology 33: 693-703

  • {199}

    SPEAKMAN, J.R., Talbot, D.A., Selman, C., Snart, S., McLaren, J.S., Redman, P., Krol, E., Jackson, D.M., Johnson,M.S. & Brand, M.D. (2004)

    Uncoupled and surviving: individual mice with high metabolism have greater mitochondrial uncoupling and live longer.
    . Aging Cell 3:87-95

    This paper was featured on the front of the journal

    It was widely reported in the popular media including articles in
    The Aberdeen Evening Express
    The Daily mail
    The Aberdeen Press and Journal
    The Glasgow herald
    The Scotsman (Front page article)
    BBC teletext
    The Sun

    The paper was featured in several science news sites on the internet including
    VPRO Dutch television Science web site
    Science Blog
    Medical news today
    Scientific approachs ? breaking news
    The paper was also subject of independent review articles in
    Nature (Nature Science update 3rd June 2004)
    Science (SAGE-KE web site lead article 28th May 2004)
    Discover magazine (

    Plus a 5 page commentary was published in Aging cell on the broad implications and importance of the work as a milestone in our understanding of links between ageing and metabolism.
    See Van Voorhies W.A. (2004) Commentary on Speakman et al : Ageing Cell 3:

    Highly cited :By february 2009 this paper was the highest cited paper ever published in the journal Aging Cell since its launch in 2002. By January 2012 it was still in this top position from a total of 705 papers ever published by this journal.

  • {198}

    SPEAKMAN, J.R. and Krol, E. (2005)

    Validation of the doubly-labelled water method in a small mammal

    Physiological and Biochemical Zoology 78: 650-667

  • {197}

    SPEAKMAN, J.R. (2004)

    Obesity: the integrated roles of environment and genetics.

    Journal of Nutrition.134: 2090-2105S

  • {196}

    Wiersma, P., Selman, C., SPEAKMAN, J.R. and Verhulst, S. (2004)

    Birds sacrifice oxidative protection for reproduction

    Proceedings of the Royal Society Series B 271: 360-365S

  • {195}

    SPEAKMAN, J.R., Krol, E. and Johnston, M.S. (2004)

    The functional significance of individual variations in BMR.

    Physiological and Biochemical Zoology 77: 900-915

    In September 2009 this paper was listed on the PBZ web site as the 4th most highly cited paper over the past 3 years.


  • {194}

    D.M. Scantlebury, Shanas, U., Kupshtein, H., SPEAKMAN, J.R. and Haim, A.(2004)

    Differential energy costs of winter acclimatized common spiny mice Acomyscahirinus from two adjacent habitats

    Comparative Physiology and Biochemistry A 137: 419-423

  • {193}

    Irwin, M.R. and SPEAKMAN, J.R. (2003)

    Azorean bats cluster as they emerge from roosts despite the lack of avian predators.

    Acta Chiropterologica

  • {192}

    Peacock WL, Krol E, Moar KM, McLaren JS, Mercer JG & SPEAKMAN, J.R. (2003)

    Photoperiodic effects on body mass, energy balance and hypothalamic gene expression in the bank vole.

    Journal of Experimental Biology 207: 165-177

  • {191}

    Butler, P.J., Green, J.A., Boyd, I.L. and SPEAKMAN, J.R. (2004)

    Measuring metabolic rate in the field: the pros and cons of the doubly-labelled water and heart rate methods

    Functional Ecology 18: 168-183

    Highly cited: By 2011 this was the 4th highest cited paper in Functional Ecology from 117 articles published in 2004

  • {190}

    SPEAKMAN, J.R., Ergon, T.E., Scantlebury, M., Reid, K.A., Cavagne, L. and Lambin, X. (2003)

    Resting metabolic rate is correlated with daily energy expenditure in free-living voles (Microtus agrestis) but reflects extrinsic rather than intrinsic limits.

    Proceedings of the National Academy of Sciences of the United States of America100: 14057-14062

  • {189}

    Hochscheid, S., Bientivegna, F. and SPEAKMAN, J.R. (2004)

    Long term cold acclimation leads to high RQ effects on oxygen consumption in loggerhead sea turtles (Caretta caretta).

    Physiologial and Biochemical Zoology 77: 209-222

  • {188}

    Krol, E., Johnson, M. S. and SPEAKMAN, J. R. (2003).

    Limits to sustained energy intake. VIII. Resting metabolic rate and organ morphology of laboratory mice lactating at thermoneutrality.

    Journal of Experimental Biology 206: 4283-4291

  • {187}

    Krol, E. and SPEAKMAN, J. R. (2003).

    Limits to sustained energy intake. VII. Milk energy output in laboratory mice at thermoneutrality

    Journal of Experimental Biology 206: 4267-4281

  • {186}

    Krol, E. and SPEAKMAN, J. R. (2003).

    Limits to sustained energy intake. VI. Energetics of lactation in laboratory mice at thermoneutrality.

    Journal of Experimental Biology 206: 4255-4266

  • {185}

    SPEAKMAN, J.R., van Acker, A., and Harper, E.J. (2003)

    Age related changes in the metabolism and body composition of three dog breeds and their relationship to life expectancy.

    Aging Cell.2: 265-279

    (see independent review of this paper in Science 12th Sept 2003)

  • {184}

    Ergon,T., SPEAKMAN, J.R., Scantlebury, M., Cavanagh, R. and Lambin, X.(2004)

    Optimal body size and energy expenditure during winter: why are voles smaller in declining populations?

    American Naturalist 163: E442-E457

  • {183}

    Hochschied, S., Bientivegna, F., and SPEAKMAN, J.R. (2003)

    The dual function of the chelonian lung: buoyancy control and oxygen storage.

    Journal of Experimental Marine Biology and Ecology 297: 123-140

  • {182}

    Schmidt, J., Andersen, N.A, SPEAKMAN, J.R. and Nicol, S.C. (2003)

    Field energetics of free-living, lactating and non-lactating echidnas (Tachyglossus aculeatus)

    Comparative Physiology and Biochemistry A 136: 903-909

  • {181}

    SPEAKMAN, J.R. and Selman, C. (2003)

    Physical activity and resting metabolic rate.

    Proceedings of the Nutrition Society 62: 621-634

  • {180}

    Korine, C., SPEAKMAN, J.R. & Arad, Z. (2004)

    Reproductive energetics of captive and free-ranging Egyptian fruit bats (Rousettus aegyptiacus)

    Ecology 85: 220-230

  • {179}

    Massemin, S., Korpim?ki, E., Zorn, T., P?yri, V. & SPEAKMAN, J.R. (2004)

    Nestling energy expenditure of Eurasian kestrels Falco tinnunculus in relation to food intake and hatchling order

    Journal of Avian Biology 3: 1-12

  • {178}

    Ward, S., SPEAKMAN, J.R. and Slater, P.J.B. (2003)

    The energy cost of song in the canary, Serinus canaria

    Animal behaviour 66: 93-902

  • {177}

    Scantlebury, M, Shanas, U., SPEAKMAN, J.R., Kupshtein, H., Afik, D. and Haim, A. (2003).

    Energetics and water economy of common spiny mice Acomys cahirinus from north and south-facing slopes of a Mediterranean valley

    Functional ecology 17: 178-185

  • {176}

    Hatch, K.A., Pinshow, B. and SPEAKMAN, J.R. (2002)

    The analysis of C-13/C-12 ratios in exhaled CO2: Its advantages and potential application to field research to infer diet, changes in diet over time, and substrate metabolism in birds

    Integrative and Comparative Biology 42: 21-33.

  • {175}

    Bunger, L., Forsting, J., McDonald, K.L., Horvat, S., Duncan, J., Hochscheid, S., Baile, C.A., Hill, W.G. and SPEAKMAN, J.R. (2002)

    Long-term divergent selection on body fatness in mice indicates a regulation system that is independent of leptin production and reception

    FASEB Journal 16:U463-477

  • {174}

    SPEAKMAN, J.R., Stubbs, R.J. and Mercer, J.G. (2002)

    Does body mass play a role in the regulation of food intake?

    Proceedings of the Nutrition Society 61: 473-487

  • {173}

    Scantlebury, M., Russell, A.F., McIlrath, G.M., SPEAKMAN, J.R.and Clutton-Brock, T.H. (2002)

    The energetics of lactation in cooperative breeding meerkatsSuricata suricatta.

    Proceedings of the Royal Society of London Series B-Biological Sciences 269: 2147-2153

    This paper was featured on the front cover of the issue.

  • {172}

    Hambly, C., Harper, E.J. and SPEAKMAN, J.R. (2002)

    The cost of flight in the zebra finch (Taenyopygia guttata) : a novel approach based on elimination of carbon-13 labelled bicarbonate.

    Journal of Comparative physiology. 172: 529-539

  • {171}

    Humphries, M.M., Thomas, D.W. & SPEAKMAN, J.R. (2002)

    Climate mediated energetic constraints on the distribution of hibernating bats.

    Nature 418: 313-316

  • {170}

    Hatch, K.A., Pinshow, B. and SPEAKMAN, J.R. (2002)

    Carbon isotope ratios in exhaled CO2 can be used to determine not just present, but also past diets in birds

    Journal of Comparative Physiology B: 172: 263-268

  • {169}

    Selman, C., Grune, T., Stolzing, A., Jakstadt, M., McLaren, J.S, & SPEAKMAN, J.R. (2002)

    The consequences of acute cold exposure on protein oxidation and proteasome activity in short-tailed field voles, Microtus agrestis

    Free Radical Biology and Medicine 33: 259-265.

  • {168}

    Selman, C., McLaren, J.S., A.R. Collins, G.G. Duthie andSPEAKMAN, J.R. (2002)

    Anti-oxidant enzyme activities, lipid peroxidation and DNA oxidative damage: the effects of short-term voluntary wheel running.

    Archives of Biochemisty and Biophysics 401: 255-261

  • {167}

    Humphries, M.M., Thomas, D.W., Hall, C.L.,SPEAKMAN, J.R.., and Kramer, D.L. (2002)

    The energetics of autumn mast hoarding in eastern chipmunks

    Oecologia 133: 30-37

  • {166}

    Hochscheid, S., Bentivegna, F. and SPEAKMAN, J.R. (2002)

    Regional blood flow in sea turtles: implications for heat exchange in an aquatic ectoterm.

    Physiological and Biochemical Zoology 75: 66-76,

  • {165}

    SPEAKMAN, J.R., Selman, C., McLaren, J.S. and Harper, J.E. (2002)

    Living fast, dying when? The links between energetics and ageing.

    Journal of Nutrition 132: 1583-1597S.

  • {164}

    Lancaster, W.C. and SPEAKMAN, J.R. (2002)

    Variations in respiratory muscle activity during echolocation when stationary in three species of bats (Microchiroptera: vespertilionidae).

    Journal of Experimental Biology 204: 4185-4193

  • {163}

    Russ, J.M., Hutson, A.M., Montgomery, W.I., Racey, P.A. andSPEAKMAN, J.R. (2001)

    The status of Nathusius' pipistrelle (Pipistrellus nathusii Keyserling & Blasius, 1839) in the British Isles

    Journal of Zoology (London) 254: 91-100.

  • {162}

    Ward, S., Moller, U., Rayner, J.M.V., Moller, U., Jackson, D.M., Biol, U., Nachtigall, W. and SPEAKMAN, J.R. (2001)

    Matabolic power, mechanical power and efficiency during wind tunnel flight in the European starling (Sturnus vulgaris).

    Journal of Experimental Biology 204: 3311-3322.

  • {161}

    .Bryce, J.M., SPEAKMAN, J.R., Johnson, P.J.and MacDonald, D.W. (2001)

    Competition between Eurasian red and introduced Eastern grey squirrels: the energetic significance of body-mass differences.

    Proceedings of the Royal Society of London Series B-Biological Sciences 268 Iss 1477: 1731-1736.

  • {160}

    Perkins, S.E. and SPEAKMAN, J.R. (2001)

    Measuring natural abundance of C-13 in respired CO2: variability and implications for non-invasive dietary analysis

    Functional Ecology 15:791-797.

  • {159}

    Selman, C., Lumsden, S., Korhonen, T., Bunger, L., Hill, W.G. andSPEAKMAN, J.R. (2001)

    Thermoregulatory responses of two mouse Mus musculus strains selectively bred for high and low food intake

    Journal of Comparative Physiology B 171: 661-668

  • {158}

    Peacock, W. and SPEAKMAN, J.R. (2001)
    Effect of high-fat diet on body mass and energy balance in the bank vole.
    Physiology and Behavior 74: 65-70.

  • {157}

    Scantlebury, D.M., Butterwick, R. and SPEAKMAN, J.R. (2001)
    Energetics and litter size variation in domestic dog Canis familiaris breeds of two sizes.
    Comparative Biochemistry and Physiology A - Molecular and Integrative Physiology129 Iss 4:

  • {156}

    Thomas, D.W., Blondel, J., Perret, P., Lambrechts, M.M. andSPEAKMAN, J.R. (2001)

    Energetic and fitness costs of mismatching resource supply and demand in seasonally breeding birds.

    Science 291 Iss 5513: 2598-2600.

  • {155}

    Jackson, D.M., Trayhurn, P. and SPEAKMAN, J.R. (2001)

    Associations between energetics and over-winter survival in the short-tailed field vole Microtus agrestis.

    Journal of Animal Ecology 70 , 633-640.

  • {154}

    Thomson, S.C., Brooke, A.P. and SPEAKMAN, J.R. (2002)
    Soaring behaviour in the Samoan flying fox, Pteropus samoensis.
    Journal of Zoology (London).256: 55-62

  • {153}

    Gremillet, D., Wanlass, S., Carss, D.N., Linton, D., Harris, M.P.,SPEAKMAN, J.R. and Le Maho, Y. (2001)

    Foraging energetics of arctic cormorants and the evolution of diving birds.

    Ecology Letters 4 Iss 3: 180-184.

     This paper was featured on the front cover of the issue.

    See also independent review of this paper in TREE (July 2001)

  • {152}

    Mercer, J.G. and SPEAKMAN, J.R. (2001)

    Hypothalamic neuropeptide mechanisms for regulating energy balance: from rodent models to human obesity.

    Neuroscience and Biobehavioral Reviews 25: 101-116.

  • {151}

    Scantlebury, D.M., Butterwick, R. and SPEAKMAN, J.R. (2001)

    Energetics and litter size variation in domestic dog Canis familiaris breeds of two sizes.

    Comparative Biochemistry and Physiology A - Molecular and Integrative Physiology129: 919-931.


    We measured resting metabolic rate ?RMR., daily energy expenditure ?DEE. and metabolisable energy intake ?MEI.
    in two breeds of dog during peak lactation to test whether litter size differences were a likely consequence of allometric
    variation in energetics. RMR of Labrador retrievers ?30 kg, n12. and miniature Schnauzers ?6 kg, n4. averaged
    3437 and 1062 kJday, respectively. DEE of Labradors ?n6. and Schnauzers ?n4. averaged 9808 and 2619 kJday,
    respectively. MEI of Labradors ?n12. was 22448 kJday and of Schnauzers ?n7. was 5382 kJday. DEE of
    Labrador pups ?2.13 kg, n19. was 974 kJday and Schnauzers ?0.89 kg, n7. were 490 kJday. Although Labradors
    had higher MEIs than Schnauzers during peak lactation, there was no difference in mass-specific energy expenditure
    between the two breeds. Hence, it is unlikely that litter size variation is a likely consequence of differences in maternal
    energy expenditure. Individual offspring were relatively more costly for mothers of the smaller breed to produce.
    Therefore, litter size variations were consistent with the expectation that smaller offspring should be more costly for
    mothers, but not that smaller mothers should per se invest more resources in reproduction. 

  • {150}

    Johnson, M.S., Thomson, S.C. and SPEAKMAN, J.R. (2001)

    Limits to sustained energy intake I. Lactation in the laboratory mouse Mus musculus.

    Journal of Experimental Biology 204: 1925-1935.


    Laboratory mice (strain MF1) were used to determine
    whether sustainable rates of energy intake are limited
    during lactation. Mice raising natural-sized litters (N=71)
    reached an asymptote in their daily food intake between
    days 13 and 16 of lactation at 23.1 g day-1 and also between
    litter sizes of 9 and 15 pups (22.8 g day-1). A second group
    of 37 females had their litter sizes manipulated at birth to
    raise more or fewer offspring than they gave birth to. When
    the litter size was increased, females did not increase their
    food intake to match their new litter size. However, when
    litter size was decreased, females decreased their
    asymptotic daily food intake during late lactation in
    relation to the extent of reduction in litter size. Therefore,
    it appeared that females were limited during late lactation
    and with large litter sizes. The milk energy exported
    amounted to 44 % of the gross energy intake, and the
    estimated daily energy expenditure was therefore
    considerably lower than the sustained energy intake
    [8.0´RMR(gross), 6.6´RMR(assimilated)], and averaged
    3.1´RMR, where RMR is resting metabolic rate. It was
    not possible to determine whether the apparent limit
    on sustained energy intake was acting centrally or
    peripherally because of the asymptotes in both food intake
    and milk energy output with increasing litter size.

  • {149}

    Johnson, M.S., Thomson, S.C. and SPEAKMAN, J.R. (2001)

    Limits to sustained energy intake II. Inter-relationships between resting metabolic rate, life-history traits and morphology in Mus musculus.

    Journal of Experimental Biology 204: 1937-1946.


    Links between resting metabolic rate (RMR) and
    reproductive output have been previously sought at both
    inter- and intraspecific levels, but have only been found in
    some interspecific studies. We aimed to examine
    correlations between RMR measured both prior to
    breeding and at peak lactation with litter size and litter
    mass in Mus musculus. By manipulating the litter size of
    some females at birth, we aimed to establish the direction
    of causality in any correlation between litter size and RMR.
    Correlations between maternal morphology and RMR,
    litter size and litter mass were also examined. Neither prebreeding
    RMR nor mass-independent pre-breeding RMR
    was correlated with litter size or litter mass. RMR at peak
    lactation, however, was positively correlated with litter size
    and negatively correlated with mean pup mass. After
    correcting for the effects of body mass, residual peak
    lactation RMR was not correlated with litter size or litter
    mass. Body size was the major morphological variable
    influencing litter mass, offspring mass and asymptotic food
    intake. Mammary tissue mass was correlated with litter
    size when only the data for mice raising unmanipulated
    litters were used. RMR at peak lactation was significantly
    related to the principal component of morphology
    dominated by carcass mass. This study confirms the
    findings of previous intraspecific and some interspecific
    studies that found no correlation between RMR and
    reproductive output after the effects of body mass had been

  • {148}

    Johnson, M.S., Thomson, S.C. and SPEAKMAN, J.R. (2001)

    Limits to sustained energy intake III. Effects of concurrent pregnancy and lactation in Mus


    Journal of Experimental Biology 204: 1947-1956.


    To determine whether mice were limited in their
    capacity to absorb energy during late lactation, we
    attempted to increase the energy burden experienced by a
    group of female mice during late lactation by mating them
    at the postpartum oestrus, hence combining the energy
    demands of pregnancy and lactation. These experimental
    mice were therefore concurrently pregnant and lactating in
    their first lactation, and were followed through a normal
    second lactation. In a control group, females also
    underwent two lactations but sequentially, with the second
    mating after the first litter had been weaned. Maternal
    mass and food intake were measured throughout the first
    lactation, second pregnancy and second lactation. Maternal
    resting metabolic rate (RMR) was measured prior to the
    first mating and then at the peak of both the first and
    second lactations. Litter size and litter mass were also
    measured throughout both lactations. In the first lactation,
    experimental mice had a lower mass-independent RMR
    (F1,88=5.15, P=0.026) and raised significantly heavier pups
    (t=2.77, d.f.=32, P=0.0093) than the control mice.
    Experimental mice delayed implantation at the start of the
    second pregnancy. The extent of the delay was positively
    related to litter size during the first lactation (F1,19=4.58,
    P=0.046) and negatively related to mean pup mass
    (F1,19=5.78, P=0.027) in the first lactation. In the second
    lactation, the experimental mice gave birth to more (t=2.75,
    d.f.=38, P=0.0092) and lighter (t=-5.01, d.f.=38, P<0.0001)
    pups than did the controls in their second lactation.
    Maternal asymptotic daily food intake of control mice in
    the second lactation was significantly higher (t=-4.39,
    d.f.=37, P=0.0001) than that of the experimental mice and
    higher than that of controls during their first lactation.
    Despite the added burden on the experimental females
    during their first lactation, there was no increase in their
    food intake, which suggested that they might be limited by
    their capacity to absorb energy. However, control females
    appeared to be capable of increasing their asymptotic food
    intake beyond the supposed limits estimated previously,
    suggesting that the previously established limit was not a
    fixed central limitation on food intake. As RMR increased
    in parallel with the increase in food intake during the
    second lactation of control mice, the sustained energy
    intake remained at around 7.0´RMR.

  • {147}

    SPEAKMAN, J.R., Gidney, A., Bett, J., Mitchell, I.P. and Johnson, M.S. (2001)

    Limits to sustained energy intake IV. Effect of variation in food quality on lactating mice Mus musculus.

    Journal of Experimental Biology 204: 1957-1965.


    Observations were made on 30 MF1 mice with their
    litters. The animals were fed either normal pelleted mouse
    food (SDS BP Nutrition Ltd) containing 13.4 kJ g-1
    digestible energy or a specially formulated diet that
    provided 25 % less digestible energy (9.75 kJ g-1) but
    equivalent amounts of protein and essential minerals and
    vitamins per gram as the normal diet. Half the animals
    were switched to the low-energy diet during early
    pregnancy and half after parturition. The food intake
    of the two groups increased enormously following
    parturition, reaching an asymptote over the last few days
    of lactation. In both groups, the asymptotic food intake
    exceeded that previously observed across 71 litters of this
    strain of mice fed the normal diet throughout pregnancy
    and lactation; the intake of the group fed the low-energy
    diet from early lactation significantly exceeded that of the
    mice switched to the low-energy diet after parturition. The
    increased intakes of the experimental groups were,
    however, insufficient to offset the lower digestible energy
    content of the food during lactation. The body mass of the
    mothers at the end of lactation did not differ between the
    two experimental groups and the controls. Offspring mass
    at weaning was inversely related to litter size, but also did
    not differ between the three groups; pup mortality did not
    differ between the experimental and control groups.
    Behavioural observations showed that during both the
    dark and light phases the general activity of the mother
    declined enormously from early pregnancy to late
    lactation. In the dark phase, the time spent in general
    activity was replaced by time spent both feeding and
    resting (suckling young), but in the light phase it was
    replaced only by feeding. At peak lactation, the mice fed
    for 30–50 % of the dark phase and for 30–40 % of the light
    phase. The data indicate that a previously observed
    asymptote in food intake during peak lactation at 23 g day-1
    is unlikely to be a limit mediated centrally by the
    alimentary tract. A higher central limit may exist, at
    26.9 g day-1, but this is unlikely to reflect the time available
    for feeding. The data are consistent with limits on
    sustainable daily energy intake being mediated by the
    performance of the mammary glands. Animals appeared to
    accommodate the demands for milk production within a
    constrained total energy budget by compensating their
    behaviour, most notably by reductions in the time spent in
    ‘general activity’.

  • {146}

    Johnson, M.S. and SPEAKMAN, J.R. (2001)

    Limits to sustained energy intake V. Effect of cold-exposure during lactation in Mus musculus.

    Journal of Experimental Biology 204: 1967-1977


    We have previously observed that female MF1 mice

    appeared to reach a limit in their food intake and milk
    production during late lactation, reaching a plateau
    between days 13 and 16 of lactation and between litter sizes
    of 9 and 15. These mice did not increase their food intake
    when forced to raise more offspring or when manipulated
    to be concurrently pregnant during late lactation, yet they
    did eat significantly more food at the peak of their second
    sequential lactation or when challenged with food of
    reduced energy content. These data suggest that apparent
    limits on sustained energy intake in this strain may not
    reflect central limitations but rather peripheral constraints
    at the mammary glands. In this study, we aimed to
    determine whether these were indeed limits by increasing
    the demands on the females during late lactation by coldexposure
    (8 °C). Females responded to this manipulation by
    significantly increasing their food intake (F1,73=77.53,
    P<0.001) above that of lactating females kept in warmer
    conditions (21 °C). In addition, there was a significant
    reduction in the number of pups raised in the cold (t=2.36,
    d.f.=18, P=0.03), with the majority of the mortality
    occurring within the first 2 days of cold-exposure. The
    mean mass of the pups raised in the cold was significantly
    lower (F1,74=13.8, P<0.001) than that of those raised in the
    warm. Despite the cold-exposure and the increased food
    intake, there was no difference in the resting metabolic
    rates of the two groups of mothers or in the lengths of their
    small intestine. The greater food intake of lactating mice
    during cold-exposure supported our previous observations
    that they were capable of eating more food than the
    previously suggested limit of 23.1 g day-1. However, the
    milk energy output of females in the cold was also
    significantly higher than in the warm (F1,15=11.99,
    P=0.003), indicating that the asymptotic food intake of
    females in the warm was not mediated by limitations in
    their milk production. Sustained energy intake in these
    mice does not appear to be centrally or peripherally
    limited. Rather, the mice may restrain their use of energy
    during their first lactation because of life-history
    consequences for future reproductive attempts.
    Key words: energetics, maximal metabolic rate, sustained

  • {145}

    Selman, C., Lumsden, S., Bunger, L., Hill, W.G. and SPEAKMAN, J.R. (2001)

    Resting metabolic rate and morphology in mice (Mus musculus) selected for high and low food intake.

    Journal of Experimental Biology 204: 777-784

  • {144}

    Schmid, J. and SPEAKMAN, J.R. (2000)

    Gender related variation in the daily energy expenditure of the grey mouse lemur (Microcebus murinus): a small primate that uses torpor.

    Journal of Comparative Physiology B – Biochemical Systemic and Environmental Physiology 170: 633-641.

  • {143}

    SPEAKMAN, J.R., Perez-Camargo, G., McCappin, T., Frankel, T., Thomson, P. and Legrand-Defretin, V. (2001)
    Validation of the doubly-labelled water technique in the domestic dog (Canis familiaris).
    British Journal of Nutrition 85: 75-87.

  • {142}

    SPEAKMAN, J.R., Booles, D. and Butterwick, R.F. (2001)
    Validation of dual energy X-ray absorptiometry (DXA) by comparison with chemical analysis of dogs and cats.
    International Journal of Obesity 25: 439-447

  • {141}

    SPEAKMAN, J.R. (2001)
    The evolution of flight and echolocation in bats: another leap in the dark.
    Mammal Review 31: 111-130.

  • {140}

    Hawkins, P.A.J., Butler, P.J., Woakes, A.J. and SPEAKMAN, J.R.(2000)
    Estimation of the rate of oxygen consumption of the common eider duck (Somateria mollissima), with some measurements of heart rate during voluntary dives.
    Journal of Experimental Biology 203: 2819-2832.

  • {139}

    Selman, C., McLaren, J.S., Himanka, M.J. and SPEAKMAN, J.R.(2000)
    Effects of long-term cold exposure on antioxidant enzyme activities in a small mammal.
    Free Radical Biology and Medicine 28: 1279-1285.

  • {138}

    Hill, R.C., Lewis, D.D., Scott, K.C., Omori, M., Jackson, M., Sundstrom, D.A., Jones, G.L., SPEAKMAN, J.R., Doyle, C.A. and Butterwick, R.F. (2001)
    Effect of increased dietary protein and decreased dietary carbohydrate on performance and body composition in racing greyhounds.
    American Journal of Veterinary Research 62: 440-447.

  • {137}

    Jackson, D.M., Hambly, C., Trayhurn, P. and SPEAKMAN, J.R.(2001)
    Can non-shivering thermogenesis in brown adipose tissue following NA injection be quantified by changes in overlying surface temperatures using infrared thermography?
    Journal of Thermal Biology 26: 85-93.

  • {136}

    Rayner, J.M.V., Viscardi, P., Ward, S., and SPEAKMAN, J.R. (2002)
    Aerodynamics and energetics of intermittent flight in birds.
    American Zoologist 41: 188-204.

  • {135}

    Ward, S., Scantlebury, D.M., Krol, E., Thomson, P.J., Sparling, C. and SPEAKMAN, J.R. (2000)
    Preparation of hydrogen from water by reduction with lithium aluminium hydride for the analysis of ? 2H by isotope ratio mass spectrometry.
    Rapid Communications in Mass Spectrometry 14: 450-453.

  • {134}

    Entwistle, A.C., Racey, P.A. and SPEAKMAN, J.R. (2000)
    Social and population structure of a gleaning bat, Plecotus auritus.
    Journal of Zoology 252: 11-17.

  • {133}

    McLean, J.A. and SPEAKMAN, J.R. (2000)
    Effects of body mass and reproduction on the basal metabolic rate of brown long-eared bats (Plecotus auritus).
    Physiological and Biochemical Zoology 73: 112-121.

  • {132}

    Redman, P., Selman, C. and SPEAKMAN, J.R. (1999)
    Male short-tailed field voles (Microtus agrestis) build better insulated nests than females.
    Journal of Comparative Physiology B ? Biochemical Systemic and Environmental Physiology 169: 581-587.

  • {131}

    SPEAKMAN, J.R. & Rydell, J. (2000)
    Avoidance behaviour of bats and moths: when is it predator defence?
    Oikos 88: 221-223.

  • {130}

    Scantlebury, D.M., Butterwick, R. and SPEAKMAN, J.R. (2000)
    Energetics of lactation in domestic dog (Canis familiaris) breeds of two sizes.
    Comparative Biochemistry and Physiology A ? Molecular and Integrative Physiology 125:197-210.

  • {129}

    Scantlebury, D.M.., Hynds, W., Booles, D. and SPEAKMAN, J.R.(2000)
    Isotope recycling in lactating dogs (Canis familiaris).
    American Journal of Physiology - Regulatory Integrative and Comparative Physiology 278: R669-R676.

  • {128}

    Krol, E. and SPEAKMAN, J.R. (1999)
    Isotope dilution spaces of mice injected simultaneously with deuterium, tritium and oxygen-18.
    Journal of Experimental Biology 202: 2839-2849.

  • {127}

    SPEAKMAN, J.R., Irwin, N.R., Tallach, N. and Stone, R.B. (1999)
    Effect of roost size on the emergence behaviour of pipistrelle bats.
    Animal Behaviour 58: 787-795.

  • {126}

    SPEAKMAN, J.R. (2000)
    The cost of living: Field metabolic rates of small mammals.
    Advances in Ecological Research 30: 177-297.

  • {125}

    SPEAKMAN, J.R. (1999)
    The evolution of flight and echolocation in pre-bats: an evaluation of the energetic efficiency of reach hunting.
    Acta Chiropterologica 1: 3-16.

  • {124}

    Thomson, S.C. and SPEAKMAN, J.R. (1999)
    Absorption of visible spectrum radiation by the wing membranes of living pteropodid bats.
    Journal of Comparative Physiology B - Biochemical Systemic and Environmental Physiology 169: 187-194

  • {123}

    McLean, J.A. and SPEAKMAN, J.R. (2000)
    Morphological changes during postnatal growth and reproduction in the brown long-eared bat Plecotus auritus: implications for wing loading and predicted flight performance.
    Journal of Natural History 34: 773-791.

  • {122}

    SPEAKMAN, J.R., Rydell, J., Webb, P.I., Hayes, J.P., Hays, G.C., Hulbert, I.A.R. and McDevitt, R.M. (2000)
    Activity patterns of insectivorous bats and birds in northern Scandinavia (69?N), during continuous midsummer daylight.
    Oikos 88: 75-86.

    See also independent reviews of this paper in
    New Scientist (Oct 1999)
    Canadian Wildlife (Dec 1999)
    Toronto Globe and Mail (Dec 1999)
    Nature: News and Views (March 2000)

  • {121}

    Ward, S., Rayner, J.M.V., Moller, U., Jackson, D.M., Nachtigall, W. and SPEAKMAN, J.R. (1999)
    Heat transfer from starlings Sturnus vulgaris during flight.
    Journal of Experimental Biology 202: 1589-1602

  • {120}

    Corp, N., Gorman, M.L. and SPEAKMAN, J.R. (1999)
    Daily energy expenditure of free-living male Wood Mice in different habitats and seasons.
    Functional Ecology 13: 585-593

  • {119}

    SPEAKMAN, J.R. and Rossi, F.P. (1999)
    No support for socio-physiological suppression effect on metabolism of paired white mice (Mus sp.).
    Functional Ecology 13: 373-382.

  • {118}

    McLean, J.A. and SPEAKMAN, J.R. (1999)
    Energy budgets of lactating and non-reproductive brown long-eared bats (Plecotus auritus) suggest females use compensation in lactation.
    Functional Ecology 13: 360-372.

  • {117}

    Swanepoel, R.E., Racey, P.A., Shore, R.F. and SPEAKMAN, J.R.(1999)
    Energetic effects of sublethal exposure to lindane on pipistrelle bats (Pipistrellus pipistrellus).
    Environmental Pollution 104: 169-175

  • {116}

    Thompson, D.R., Bearhop, S., SPEAKMAN, J.R. and Furness, R.W. (1998)
    Feathers as a means of monitoring mercury in seabirds: insights from stable isotope analysis.
    Environmental Pollution 101: 193-200.

  • {115}

    SPEAKMAN, J.R. and Tallach, N. (1998)
    Do emerging pipistrelle bats lose control of their timing due to 'crowd pressure'?
    Journal of Zoology 246: 445-448.

  • {114}

    Jenkins, E.V., Laine, T., Morgan, S.E., Cole, K.R. and SPEAKMAN, J.R. (1998)
    Roost selection in the pipistrelle bat, Pipistrellus pipistrellus(Chiroptera: Vespertilionidae), in northeast Scotland.
    Animal Behaviour 56: 909-917.

  • {113}

    SPEAKMAN, J.R. and Rowland, A. (1999)
    Preparing for inactivity: how insectivorous bats deposit a fat store for hibernation.
    Proceedings of the Nutrition Society 58: 123-131

  • {112}

    Ward, S.M., Moller, U., Rayner, J.M.V., Jackson, D.M., Nachtigall, W. and SPEAKMAN, J.R. (1998).
    Power requirement for Starling flight in a wind tunnel.
    Biologia e Conservazione della Fauna 102: 335-339.

  • {111}

    SPEAKMAN, J.R. and Ward, S. (1998)
    Infrared thermography : principles and applications.
    Zoology - Analysis of Complex Systems. 101: 224-232.

  • {110}

    SPEAKMAN, J.R. (1998)
    The history and theory of the doubly labeled water technique.
    American Journal of Clinical Nutrition. 68: 932S-938S.

  • {109}

    Gorman, M.L., Mills, M.G., Raath, J.P.and SPEAKMAN, J.R. (1998)
    High hunting costs make African wild dogs vulnerable
    to kleptoparasitism by hyenas.
    Nature 391: 479-481.

    This paper was featured on the front cover of the issue.

    See also independent reviews of this paper in

    • Science (Jan 31 1998)
    • Washington post (Jan 31 1998)
    • The Economist (Feb 1998)
      This paper was also the subject of radio and TV broadcasts
    • Science Now (Feb 1998)
    • Discovery channel Science program (March 1998).

    See also article on this paper in the Encyclopedia Brittanica year book 1999

  • {108}

    Entwistle, A.C., Racey, P.A. and SPEAKMAN, J.R. (1998)
    The reproductive cycle and determination of sexual maturity in male brown long-eared bats, Plecotus auritus (Chiroptera: Vespertilionidae).
    Journal of Zoology 244: 63-70.

  • {107}

    Thomson, S.C., Brooke, A.P. and SPEAKMAN, J.R. (1998)
    Diurnal activity in the Samoan flying fox, Pteropus samoensis.
    Philosophical Transactions of the Royal Society of London Series B - Biological Sciences 353: 1595-1606.

    See also independent review of this paper in Nature online at

  • {106}

    SPEAKMAN, J.R., Hays, G.C. and Lindblad, E. (1998)
    Thermal conductivity of sand and its effect on the temperature of loggerhead sea turtle (Caretta caretta) nests.
    Journal of the Marine Biological Association of the United Kingdom 78: 1337-1352.

  • {105}

    SPEAKMAN, J.R. and Thomson, S.C. (1997)
    Validation of the labeled bicarbonate technique for measurement of short term energy expenditure in the mouse.
    Zeitschrift Fur Ernahrungswissenschaft 36: 273-277.

  • {104}

    SPEAKMAN, J.R. (1997)
    Factors influencing daily energy expenditure of small mammals.
    Proceedings of the Nutrition Society 56: 1119-1136.

  • {103}

    Corp, N., Gorman, M.L. and SPEAKMAN, J.R. (1997)
    Apparent absorption efficiency and gut morphometry of wood mice,Apodemus sylvaticus, from two distinct populations with different diets.
    Physiological Zoology 70: 610-614.

  • {102}

    McLean, J.A. and SPEAKMAN, J.R. (1997)
    Non-nutritional maternal support in the brown long-eared bat.
    Animal Behaviour 54: 1193-1204.

  • {101}

    SPEAKMAN, J.R. and Banks, D. (1998)
    The function of flight formations in Greylag Geese Anser anser;energy saving or orientation?
    Ibis 140: 280-287.

    See also independent reviews of this paper in
    BBC Wildlife magazine March 1999
    Nature Australia November 1999

  • {100}

    Corp, N., Gorman, M.L. and SPEAKMAN, J.R. (1997)
    Seasonal variation in the resting metabolic rate of male wood miceApodemus sylvaticus from two contrasting habitats 15 km apart.
    Journal of Comparative Physiology B - Biochemical Systemic and Environmental Physiology 167: 229-239.

  • {99}

    Webb, P.I., SPEAKMAN, J.R. and Racey, P.A. (1996)
    Population dynamics of a maternity colony of the pipistrelle bat (Pipistrellus pipistrellus) in north-east Scotland.
    Journal of Zoology 240: 777-780.

  • {98}

    Corp, N., Gorman, M.L. and SPEAKMAN, J.R. (1997)
    Ranging behaviour and time budgets of male wood mice Apodemus sylvaticus in different habitats and seasons.
    Oecologia 109: 242-250.

  • {97}

    McLean, J.A. and SPEAKMAN, J.R. (1996)
    Suckling behaviour in the brown long-eared bat (Plecotus auritus).
    Journal of Zoology 239: 411-416.

  • {96}

    Lancaster, W.C., Thomson, S.C. and SPEAKMAN, J.R. (1997)
    Wing temperature in flying bats measured by infrared thermography.
    Journal of Thermal Biology 22: 109-116.

  • {95}

    Entwistle, A.C., Racey, P.A. and SPEAKMAN, J.R. (1997)
    Roost selection by the brown long-eared bat (Plecotus auritus).
    Journal of Applied Ecology 34: 399-408.

  • {94}

    Entwistle, A.C., Racey, P.A. and SPEAKMAN, J.R. (1996)
    Habitat exploitation by a gleaning bat, Plecotus auritus.
    Philosophical Transactions of the Royal Society of London Series B - Biological Sciences 351: 921-931.

  • {93}

    McDevitt, R.M. and SPEAKMAN, J.R. (1996)
    Summer acclimatization in the short-tailed field vole, Microtus agrestis.
    Journal of Comparative Physiology B - Biochemical Systemic and Environmental Physiology 166: 286-293.

  • {92}

    Webb, P.I., SPEAKMAN, J.R. and Racey, P.A. (1996)
    How hot is a hibernaculum? A review of the temperatures at which bats hibernate.
    Canadian Journal of Zoology - Revue Canadienne de Zoologie 74: 761-765.

  • {91}

    Arnould, J.P.Y., Boyd, I.L. and SPEAKMAN, J.R. (1996)
    The relationship between foraging behaviour and energy expenditure in Antarctic fur seals.
    Journal of Zoology 239: 769-782.

  • {90}

    Hays, G.C., MacKay, A., Adams, C.R., Mortimer, J.A., SPEAKMAN, J.R. and Boerema, M. (1995)
    Nest-site selection by sea-turtles.
    Journal of the Marine Biological Association of the United Kingdom 75: 667-674.

  • {89}

    SPEAKMAN, J.R. and McQueenie, J. (1996)
    Limits to sustained metabolic rate: The link between food intake, basal metabolic rate, and morphology in reproducing mice, Mus musculus.
    Physiological Zoology 69: 746-769.

  • {88}

    McLean, J.A. and SPEAKMAN, J.R. (1995)
    Elimination rate of Zn-65 as a measure of food-intake - a validation-study in the mouse (Mus sp.).
    Journal of Applied Physiology 79: 1361-1368

  • {87}

    Cleaver, J.E., SPEAKMAN, J.R. and Volpe, J.P.G. (1995)
    Nucleotide excision repair: Variations associated with cancer development and speciation.
    Cancer Surveys 25: 125-142.

  • {86}

    SPEAKMAN, J.R. (1995)
    Energetics and the evolution of body size in small terrestrial mammals.
    Symposia of the Zoological Society of London. 69: 63-81.

  • {85}

    Arnould, J.P.Y., Boyd, I.L. and SPEAKMAN, J.R. (1996)
    Measuring the body composition of Antarctic fur seals (Arctocephalus gazella): Validation of hydrogen isotope dilution.
    Physiological Zoology 69: 93-116.

  • {84}

    Delahay, R.J., SPEAKMAN, J.R. and Moss, R. (1995)
    The energetic consequences of parasitism - effects of a developing infection of Trichostrongylus-tenuis (Nematoda) on red grouse (Lagopus-lagopus-scoticus) energy-balance, body-weight and condition.
    Parasitology 110: 473-482.

  • {83}

    SPEAKMAN, J.R. (1995)
    Estimation of precision in DLW studies using the two-point methodology.
    Obesity Research 3 (Supplement 1: 'Recent advances in the doubly labeled water technique' Eds. SPEAKMAN, J.R. and Roberts, S.B.)

  • {82}

    Haim, A., McDevitt, R.M. and SPEAKMAN, J.R. (1995)
    Daily variations in the response of wood mice Apodemus-sylvaticusto noradrenaline.
    Journal of Experimental Biology 198: 561-565.

  • {81}

    Haim, A., McDevitt, R.M. and SPEAKMAN, J.R. (1995)
    Thermoregulatory responses to manipulations of photoperiod in wood mice Apodemus-sylvaticus from high-latitudes (57 oN).
    Journal of Thermal Biology 20: 437-443.

  • {80}

    Hays, G.C., Adams, C.R., Mortimer, J.A. and SPEAKMAN, J.R.(1995)
    Inter-beach and intra-beach thermal variation for green turtle nests on Ascension Island, South-Atlantic.
    Journal of the Marine Biological Association of the United Kingdom75: 405-411.

  • {79}

    Rydell, J. and SPEAKMAN, J.R. (1995)
    Evolution of nocturnality in bats - potential competitors and predators during their early history.
    Biological Journal of the Linnean Society 54: 183-191.

    See also independent review of this paper in BBC Wildlife (May 1995)

  • {78}

    Bevan, R.M., SPEAKMAN, J.R. and Butler, P.J. (1995)
    Daily energy-expenditure of tufted ducks - a comparison between indirect calorimetry, doubly labeled water and heart-rate.
    Functional Ecology 9: 40-47.

  • {77}

    SPEAKMAN, J.R. and Thomson, S.C. (1994)
    Flight capabilities of Archaeopteryx.
    Nature 370 Iss 6490: 514.

    See also independent review of this paper in Earth Magazine (April 1995)

  • {76}

    SPEAKMAN, J.R., Hays, G.C. and Webb, P.I. (1994)
    Is hyperthermia a constraint on the diurnal activity of bats?
    Journal of Theoretical Biology 171 Iss 3: 325-339.

  • {75}
    McDevitt, R.M. and SPEAKMAN, J.R. (1994)
    Central limits to sustainable metabolic-rate have no role in cold-acclimation of the short-tailed field vole (Microtus-agrestis).
    Physiological Zoology 67 Iss 5: 1117-1139.
  • {74}

    McDevitt, R.M. and SPEAKMAN, J.R. (1994)
    Central limits to sustainable metabolic-rate have no role in cold-acclimation of the short-tailed field vole (Microtus-agrestis).
    Physiological Zoology 67: 1117-1139.

  • {73}

    SPEAKMAN, J.R., Stone, R.E. and Kerslake, J.E. (1995)
    Temporal patterns in the emergence behavior of pipistrelle bats,Pipistrellus-pipistrellus, from maternity colonies are consistent with an antipredator response.
    Animal Behaviour 50: 1147-1156

  • {72}

    SPEAKMAN, J.R. (1995)
    Chiropteran nocturnality.
    Symposium of the Zoological Society of London 67: 187-201

  • {71}

    Cutts, C.J. and SPEAKMAN, J.R. (1994)
    Energy savings in formation flight of pink-footed geese.
    Journal of Experimental Biology 189: 251-261.

  • {70}

    Webb, P.I., SPEAKMAN, J.R. and Racey, P.A. (1995)
    Evaporative water-loss in 2 sympatric species of vespertilionid bat,Plecotus-auritus and Myotis-daubentoni ? relation to foraging mode and implications for roost site selection.
    Journal of Zoology 235: 269-278.

  • {69}

    McDevitt, R.M. and SPEAKMAN, J.R. (1994)
    Long photophase is not a sufficient stimulus to reduce thermogenic capacity in winter-acclimatized short-tailed field voles (Microtus-agrestis) during long-term cold-acclimation.
    Journal of Comparative Physiology B - Biochemical Systemic and Environmental Physiology 164: 159-164.

  • {68}

    Entwistle, A.C., SPEAKMAN, J.R. and Racey, P.A. (1994)
    Effect of using the doubly labeled water technique on long-term recapture in the brown long-eared bat (Plecotus-auritus).
    Canadian Journal of Zoology 72 Iss 4: 783-785.

  • {67}

    Poppitt, S.D., SPEAKMAN, J.R. and Racey, P.A. (1994)
    Energetics of reproduction in the lesser hedgehog tenrec, Echinops telfairi (Martin).
    Physiological Zoology 67 Iss 4: 976-994.

  • {66}

    SPEAKMAN, J.R., McDevitt, R.M. and Cole, K.R. (1993)
    Measurement of basal metabolic rates - don't lose sight of reality in the quest for comparability.
    Physiological Zoology 66 Iss 6: 1045-1049.

  • {65}

    Gauthier, M., Thomas, D.W., SPEAKMAN, J.R. and LaPierre (1994)
    The costs and benefits of nest reuse by cliff martins. (In French English Abstract.)

  • {64}

    Blake, D., Hutson, A.M., Racey, P.A., Rydell, J. and SPEAKMAN, J.R. (1994)
    Use of lamplit roads by foraging bats in southern England.
    Journal of Zoology 234: 453-462.

  • {63}

    SPEAKMAN, J.R., Racey, P.A., Haim, A., Webb, P.I., Ellison, G.T.H. and Skinner, J.D. (1994)
    Interindividual and intraindividual variation in daily energy-expenditure of the pouched mouse (Saccostomus-campestris).
    Functional Ecology 8 Iss 3: 336-342.

  • {62}

    Webb, P.I., SPEAKMAN, J.R. and Racey, P.A. (1994)
    Postprandial urine loss and its relation to ecology in brown long-eared (Plecotus-auritus) and Daubentons (Myotis-daubentoni) bats (Chiroptera, Vespertilionidae).
    Journal of Zoology 233: 165-173.

  • {61}

    SPEAKMAN, J.R. (1993)
    How should we calculate CO2 production in doubly labeled water studies of animals?
    Functional Ecology 7: 746-750.

  • {60}

    Stephenson, P.J., SPEAKMAN, J.R. and Racey, P.A. (1994)
    Field metabolic-rate in 2 species of shrew-tenrec, Microgal- dobsoniand M-talazaci.
    Comparative Biochemistry and Physiology A - Physiology 107 Iss 2: 283-287.

  • {59}

    Webb, P.I., SPEAKMAN, J.R. and Racey, P.A. (1993)
    The implication of small reductions in body-temperature for radiant and convective heat-loss in resting endothermic brown long-eared bats (Plecotus auritus).Journal of Thermal Biology 18: 131-135.

  • {58}

    Siegert, K.J., SPEAKMAN, J.R. and Reynolds, S.E. (1993)
    Carbohydrate and lipid-metabolism during the last larval molt of the tobacco hornworm, Manduca-sexta.
    Physiological Entomology 18 Iss 4: 404-408.

  • {57}

    Hays, G.C., Adams, C.R. and SPEAKMAN, J.R. (1993)
    Reproductive investment by green turtles nesting on Ascension-Island.
    Canadian Journal of Zoology 71: 1098-1103.

  • {56}

    SPEAKMAN, J.R. and Hays, G.C. (1992)
    Albedo and transmittance of short-wave-radiation for bat wings.
    Journal of Thermal Biology 17: 317-321.

  • {55}

    SPEAKMAN, J.R. and Webb, P.I. (1993)
    Taxonomy, status and distribution of the Azorean bat (Nyctalus-azoreum).
    Journal of Zoology 231: 27-38.

  • {54}

    SPEAKMAN, J.R., Nair, K.S. and Goran, M.I. (1993)
    Revised equations for calculating CO2 production from doubly labeled water in humans.
    American Journal of Physiology 264: E912-E917.

  • {53}

    Rydell, J., Strann, K.B. and SPEAKMAN, J.R. (1994)
    First record of breeding bats above the Arctic-circle - northern bats at 68-70oN in Norway.
    Journal of Zoology 233: 335-339.

  • {52}

    Poppitt, S.D., SPEAKMAN, J.R. and Racey, P.A. (1993)
    The energetics of reproduction in the common shrew (Sorex-araneus) - a comparison of indirect calorimetry and the doubly labeled water method.
    Physiological Zoology 66: 964-982.

  • {51}

    SPEAKMAN, J.R. and Bryant, D.M. (1993)
    The searching speeds of foraging shorebirds - Redshank (Tringa-totanus) and Oystercatcher (Haematopus-ostralegus).
    American Naturalist 142: 296-319.

  • {50}

    Webb, P.I., SPEAKMAN, J.R. and Racey, P.A. (1993)
    Defecation, apparent absorption efficiency, and the importance of water obtained in the food for water-balance in captive brown long-eared (Plecotus-auritus) and Daubentons (Myotis-daubentoni) bats.
    Journal of Zoology 230: 619-628.

  • {49}

    Brown, R.P., Perez-Mellado, V., Diego-Rasilla, J., Garcia, J.A., Naranjo, A. and SPEAKMAN, J.R. (1992)
    Individual and population energetics of a lizard on a Mediterranean islet.
    Oecologia 91: 500-504.

  • {48}

    SPEAKMAN, J.R. (1993)
    Flight capabilities in Archaeopteryx.
    Evolution 47: 336-340.

  • {47}

    SPEAKMAN, J.R. (1993)
    The evolution of echolocation for predation.
    Symposia of the Zoological Society of London 65: 39-63.

  • {46}

    SPEAKMAN, J.R., Lumsden, L.F. and Hays, G.C. (1994)
    Predation rates on bats released to fly during daylight in south-eastern Australia.
    Journal of Zoology 233: 318-321.

  • {45}

    Hayes, J.P., SPEAKMAN, J.R. and Racey, P.A. (1992)
    The contributions of local heating and reducing exposed surface-area to the energetic benefits of huddling by short-tailed field voles (Microtus-agrestis).
    Physiological Zoology. 65 Iss 4: 742-762

  • {44}

    Hays, G.C., SPEAKMAN, J.R. and Webb, P.I (1992)
    Why do brown long-eared bats (Plecotus-auritus) fly in winter?
    Physiological Zoology 65: 554-567

  • {43}

    Hays, G.C. and SPEAKMAN, J.R. (1993)
    Nest placement by loggerhead turtles, Caretta-caretta.
    Animal Behaviour 45 Iss 1: 47-53.

  • {42}

    Webb, P.I., SPEAKMAN, J.R. and Racey, P.A. (1992)
    Oxygen-consumption and evaporation during parturition in a vespertilionid bat (Pipistrellus-pipistrellus).
    Journal of Reproduction and Fertility 94: 525-528.

  • {41}

    Brown, R.B., Thorpe, R.S. and SPEAKMAN, J.R. (1992)
    Comparisons of body size, field energetics, and water flux among populations of the skink Chalcides-sexlineatus.
    Canadian Journal of Zoology 70: 1001-1006.

  • {40}

    Hayes, J.P., SPEAKMAN, J.R. and Racey, P.A. (1992)
    Sampling bias in respirometry.
    Physiological Zoology 65: 604-619.

  • {39}

    SPEAKMAN, J.R., Racey, P.A., Haim, A., Webb, P.I., Ellison, G.T.H. and Skinner, J.D. (1992)
    Daily energy-expenditure in the pouched mouse (Saccostomus-campestris Peters 1896).
    Israel Journal of Zoology 38: 341-351.

  • {38}

    Hays, G.C., SPEAKMAN, J.R. and Hayes, J.P. (1992)
    The pattern of emergence by loggerhead turtle (Caretta-caretta) hatchlings on Cephalonia, Greece.
    Herpetologica 48: 396-401.

  • {37}

    SPEAKMAN, J.R. (1992)
    Evolution of animal body size - a cautionary note on assessments of the role of energetics.
    Functional Ecology 6 Iss 4: 495-496.

    & SPEAKMAN, J.R. (1993)
    Evolution of body size - predictions from energetics.
    Functional Ecology 7 Iss 1: 134.

    These two refereed Forum articles appeared as part of a debate on the role of energetics as a factor influencing evolution of body size, principally between myself, J.S. Millar and G.J. Hickling.

  • {36}

    Hays, G.C., Webb, P.I. and SPEAKMAN, J.R. (1992)
    Arrhythmic breathing in torpid pipistrelle bats, Pipistrellus-pipistrellus).
    Respiration Physiology 85 Iss 2: 185-192.

  • {35}

    Hays, G.C. and SPEAKMAN, J.R. (1992)
    Clutch size for Mediterranean loggerhead turtles (Caretta-caretta).
    Journal of Zoology 226: 321-327.

  • {34}

    Webb, P.I., Hays, G.C., SPEAKMAN, J.R. and Racey, P.A. (1992)
    The functional-significance of ventilation frequency, and its relationship to oxygen-demand in the resting brown long-eared bat,Plecotus-auritus.
    Journal of Comparative Physiology B - Biochemical Systemic and Environmental Physiology 162: 144-147.

  • {33}

    Webb, P.I., SPEAKMAN, J.R. and Racey, P.A. (1992)
    Inter individual and intraindividual variation in wing loading and body-mass in female pipistrelle bats - theoretical implications for flight performance.
    Journal of Zoology 228: 669-673.

  • {32}

    SPEAKMAN, J.R., Webb, P.I. and Racey, P.A. (1991)

    Effects of disturbance on the energy-expenditure of hibernating bats.

    Journal of Applied Ecology 28: 1087-1104.

    see also independent reviews of this paper in
    British Wildlife (July 1991)
    The Economist (February 1993)
    this paper was featured on the Canadian Science program "Quirks and Quarks", broadcast across the US and Canada, Nov 1993.

  • {31}

    SPEAKMAN, J.R., Bullock, D.J., Eales, L.A. and Racey, P.A. (1992)

    A problem defining temporal pattern in animal behaviour - clustering in the emergence behaviour of bats from maternity roosts.

    Animal Behaviour 43: 491-500.

    see also "Defining temporal pattern in the emergence behaviour of bats from maternity roosts: some pitfalls in analysis and how to overcome them." SPEAKMAN, J.R. In Bat Research News (1993).

  • {30}

    SPEAKMAN, J.R. and Racey, P.A. (1991)
    No cost of echolocation for bats in flight.
    Nature 350 Iss 6317: 421-423.

    This paper was featured on the front cover of the issue and was the subject of a 'News and Views' article in the same issue. see also independent reviews of this paper in

    • BBC Wildlife magazine (June 1991)
    • Science News (July 1991)
    • The Lisbon Times (June 1991)
    • The European (June 1991)

    this paper was also the subject of several radio and TV broadcasts

    • Tomorrow's World (BBC 1, May 1991)
    • Radio 4 Science Now (May 1991)
    • BBC World Service
    • (syndicated worldwide: June 1991)

    see also article on this work in the McGraw-Hill Yearbook of Science and Technology (1994).

  • {29}

    SPEAKMAN, J.R., Racey, P.A., Catto, C.M.C., Webb, P.I., Swift, S.M. and Burnett, A.M. (1991)

    Minimum summer population and densities of bats in N.E. Scotland near the northern borders of their distributions.

    Journal of Zoology 225: 327-345.

  • {28}

    SPEAKMAN, J.R. (1991)

    Why do insectivorous bats in Britain not fly in daylight more frequently?

    Functional Ecology 5: 518-524.

  • {27}

    SPEAKMAN, J.R. (1991)

    The impact of predation by birds on bat populations in the British Isles.

    Mammal Review 21: 123-142

  • {26}

    SPEAKMAN, J.R., Racey, P.A. and Burnett, A.M. (1991)

    Metabolic and behavioral consequences of the procedures of the doubly labeled water technique on white (MF1) mice.

    Journal of Experimental Biology 157: 123-132.

  • {25}

    SPEAKMAN, J.R., Racey, P.A., Hutson, A.M., Webb, P.I. and Burnett, A.M. (1991)

    Status of Nathusius pipistrelle (Pipistrellus-nathusii) in Britain.

    Journal of Zoology 225: 685-690.

  • {24}

    SPEAKMAN, J.R. (1990)

    Principles, problems and a paradox with the measurement of energy-expenditure of free-living subjects using doubly-labeled water.

    Statistics in Medicine 9: 1365-1380.

  • {23}

    Hays, G.C. and SPEAKMAN, J.R. (1991)

    Reproductive investment and optimal clutch size of loggerhead sea-turtles (Caretta-caretta).

    Journal of Animal Ecology 60: 455-462.

  • {22}

    Haim, A., Racey, P.A., SPEAKMAN, J.R., Ellison, G.T.H. and Skinner, J.D. (1991)

    Seasonal acclimatization and thermoregulation in the pouched mouse Saccostomus-campestris.

    Journal of Thermal Biology 16: 13-17.

  • {21}

    SPEAKMAN, J.R. (1990)

    On Blum’s 4-dimensional geometric explanation for the 0.75 exponent in metabolic allometry.

    Journal of Theoretical Biology 144: 139-141.

  • {20}

    Hays, G.C., Webb, P.I., French, J. and SPEAKMAN, J.R. (1990)

    Doppler radar - a non invasive technique for measuring ventilation rate in resting bats.

    Journal of Experimental Biology 150: 443-447.

  • {19}

    SPEAKMAN, J.R., Nagy, K.A., Masman, D., Mook, W.G., Poppitt, S.D., Strathearn, G.E. and Racey, P.A. (1990)

    Interlaboratory comparison of different analytical techniques for the determination of O-18 abundance.

    Analytical Chemistry 62: 703-708.

  • {18}

    SPEAKMAN, J.R. (1990)

    The function of daylight flying in British bats.

    Journal of Zoology 220: 101-113.

  • {17}

    Hughes, P.M., SPEAKMAN, J.R., Jones, G. and Racey, P.A. (1989)

    Suckling behaviour in the pipistrelle bat (Pipistrellus-pipistrellus).

    Journal of Zoology 219: 665-670.

  • {16}

    SPEAKMAN, J.R. and Racey, P.A. (1989b)

    Doubly labelled water - errors in the evaluation of oxygen isotope turnover due to temporal variation in CO2 production do not always covary with dilution space estimate.

    Journal of Theoretical Biology 141: 547-556.

  • {15}

    SPEAKMAN, J.R., Anderson, M.E. and Racey, P.A. (1989)

    The energy-cost of echolocation in pipistrelle bats (Pipistrellus-pipistrellus).

    Journal of Comparative Physiology A 165: 679-685.

    This paper was featured on the front cover of the journal

  • {14}

    SPEAKMAN, J.R. and Racey, P.A. (1989)

    Hibernal ecology of the pipistrelle bat – energy-expenditure, water requirements and mass loss, implications for survival and the function of winter emergence flights.

    Journal of Animal Ecology 58: 797-813.

  • {13}

    SPEAKMAN, J.R. and Racey, P.A. (1988c)

    Validation of the doubly labeled water technique in small insectivorous bats by comparison with indirect calorimetry.

    Physiological Zoology 61: 514-526.

  • {12}

    SPEAKMAN, J.R. and Racey, P.A. (1988b)

    The doubly-labeled water technique for measurement of energy-expenditure in free-living animals.

    Science Progress 72: 227-237.

  • {11}

    SPEAKMAN, J.R. and Racey, P.A. (1988a)

    Consequences of non steady-state CO2 production for accuracy of the doubly labeled water technique – the importance of recapture interval.

    Comparative Biochemistry and Physiology A 90: 337-340.

  • {10}

    SPEAKMAN, J.R. (1988)

    Position of the pinnae and thermoregulatory status in brown long-eared bats (Plecotus-auritus).

    Journal of Thermal Biology 13: 25-29

  • {9}

    Racey, P.A., SPEAKMAN, J.R. and Swift, S.M. (1987)

    Reproductive adaptations of heterothermic bats at the northern borders of their distributions.

    South African Journal of Science 83: 635-638.

  • {8}

    Racey, P.A. and SPEAKMAN, J.R. (1987)

    The energy costs of pregnancy and lactation in heterothermic bats.

    Symposia of the Zoological Society of London 54: 107-125.

  • {7}

    SPEAKMAN, J.R. (1987b)

    Apparent absorption efficiencies for redshank (Tringa totanus L) and oystercatcher (Haematopus-ostralegus L) – implications for the predictions of optimal foraging models.

    American Naturalist 130: 677-691.

  • {6}

    SPEAKMAN, J.R. (1987a)

    Calculation of CO2 production in doubly-labeled water studies.

    Journal of Theoretical Biology 126: 101-104.

  • {5}

    SPEAKMAN, J.R. and Racey, P.A. (1987b)

    The equilibrium concentration of O-18 in body-water - implications for the accuracy of the doubly-labeled water technique and a potential new method of measuring RQ in free-living animals.

    Journal of Theoretical Biology 127: 79-95.

  • {4}

    Davidson, N.C., Townshend, D.J., Pienkowski, M.W. and SPEAKMAN, J.R. (1986)
    Why do curlews Numenius have decurved bills?
    Bird Study 33: 61-69.

  • {3}
    SPEAKMAN, J.R. and Racey, P.A. (1986b)
    The influence of body condition on sexual development of male brown long-eared bats (Plecotus-auritus) in the wild.
    Journal of Zoology (London) 210: 515-525.
  • {2}
    SPEAKMAN, J.R. and Racey, P.A. (1986a)
    Measurement of CO2 production by the doubly labeled water technique.
    Journal of Applied Physiology 61 Iss 3: 1200-1202.
  • {1}

    SPEAKMAN, J.R. (1986)
    The optimum search speed of terrestrial predators when feeding on sedentary prey - a predictive model.
    Journal of Theoretical Biology 122: 401-407.

Books and Edited Volumes

  1. J.R. SPEAKMAN and S.B. Roberts (1995)

    Recent advances in the doubly-labelled water technique.

    Proceedings of a workshop held at Anaheim (April 1994)

    Supplement to Obesity Research.

  2. J.R. SPEAKMAN(1997)

    Doubly-labelled water: Theory and practice.

    Kluwer Academic Publishers, New York

    (now Springer)

    ISBN: 0 412 63780 4

    416 pp


    Part One: Introduction to energetics. What is energetics? Methods for studying energetics. Part Two: Doubly-labelled water: Theory. Biochemical basis of the technique. Major fluxes resulting in the turnover of labels in the body. Assumptions in the model. Other issues. Validation studies in comparison to indirect calorimetry. Part Three: Doubly-labelled water: Practice. Is DLW the best technique to use? Is DLW appropriate for the animal or situation that I wish to study? Legal aspects of applying doubly-labelled water. What effect will DLW have on the subject animal? Calculating the dose to use. Buying isotopes. Field protocols. Laboratory protocols. Calculating CO2 production and energy expenditure. Part Four: Review of estimates of energy expenditure derived from studies using doubly-labelled water: Daily energy demands of wild animals. Specific applications (costs of activity). Domestic animals. Humans. Writing up doubly-labelled water studies.

    Publisher summary

    Divided into three parts, Doubly Labelled Water presents a clear and accessible account of this technique. Part One presents a general introduction to the study of animal energetics: Part Two discusses the theory behind use of doubled labellled water and Part Three evaluates the practical aspects of its use and the methodlologies required for its application.



    5.0 out of 5 stars Essential April 9, 2001
    This book really stands out, more than anything else in the last 50 years. Using doubly labelled water is complex and can be confusing, but Speakman does a great job in all three sections of his book. First, he presents the general introdution to animal energetics, then he talks about the theroy of doublly labelled water, and finally he presents useful information about the practice of both using and writing about doubly labeled water in scientific papers. These book is essential reading for graduate students and professors looking for a guide in getting started with doubly labelled water

    Robert R Harris

    Journal of Animal ecology (2002) 70: 538

    J. R. Speakman (1997) Doubly-Labelled Water – Theory and Practice. Kluwer Academic, Dordrecht. £154. ISBN: 0-412-63780-4.

    The ‘non-invasive’ doubly-labelled water (DLW) technique for determining metabolic rate measures the difference in the turnover rates of the isotopes 18O and 2H (deuterium), following their injection as 2H218O. The loss of 2H is used to calculate the 18O loss as water and, by subtracting this from the total 18O loss, the 18O loss in C18O2 production can be obtained. This can be converted to energy units. DLW techniques have been used mainly in mammals and birds, although some work has been carried out on reptiles and insects. When 18O loss as water is high – for example, due to large integumental water fluxes in aquatic ectotherms – the technique is not unsuitable. However, even in endotherms, DLW can give estimates of metabolic rate different from those obtained by more conventional methods under similar conditions.

    In the preface, Dr Speakman suggests that readers with a knowledge of animal energetics could skip Part 1 and proceed directly to Parts 2 and 3, which give the theoretical basis and practice of DLW, including detailed protocols and methods of calculation. In fact, I found Part 1 a useful synopsis of the area and background for what followed. Clear models of many underlying concepts of DLW, such as dilution, washout kinetics and compartmental analysis, are presented. However, I felt that occasionally the analogies were a little overdone, and the figure legends could have been more concise, as they seem to repeat the text in many places.

    In the Foreword, Professor Nagy tantalizingly refers to discoveries of wide differences and variations in energy expenditure which have been made using the DLW technique. Although descriptions of the fundamental experiments establishing the method are given, I did not get a feel for the real gains in our knowledge made using DLW specifically. One or two case studies, to illustrate how it has changed our ideas about animal energetics, would have helped the reader. Researchers wishing to measure the metabolic rate of animals in the field should find much useful information here, including addresses of laboratories currently using the method, suppliers of equipment and an extensive Reference section.

  3. J.R. SPEAKMAN (Editor) (2001)

    Body composition analysis in animals:

    A handbook of non-destructive methods.

    Cambridge University Press.

    ISBN: 9780521081405

    260 pp

    Publishers summary

    Bringing together a mix of traditional and well established analytical methods with more modern techniques, Body Composition Analysis of Animals provides a theoretical overview of different methodologies combined with practical advice on the use of these techniques. Methods covered include the use of destructive methods of analysis, body condition indices, isotope and gas dilution methods, total body electrical conductivity, bio-impedance analysis, ultrasound scanning, and dual energy x-ray absorptiometry.

    • Practical advice from experienced researchers
    • Information on a range of methods brought together in a single volume
    • Provides information on how different methods actually work - information that is seldom covered in primary scientific papers


    "This book has definitely fulfilled its stated goal of providing a useful summary of noninvasive methods of body composition analysis for animal sciences... This book has the potential to become a standard reference for anyone studying body composition, be it animal or human." Kenneth J. Ellis, Quarterly Review of Biology

    "Highly informative compilation containing theoretical and practical considerations of eight procedures that have been employed successfully to examine the body composition of animals... Overall, this book is user friendly, with subject and taxon indices that are thoughtfully compiled to facilitate navigation through the book and a subject index that is comprehensive... It makes for fascinating reading and should facilitate the planning of research programs, saving laborious hours of background reading and providing the wherewithal to carry out body composition analysis, both efficiently and successfully." S. James Reynolds, Ecology

    "John Speakman has brought together a group of authors who blend traditional, established, analytical methods with more cutting edge techniques and methodologies.... this book is an excellent resource and it should become a standard reference text for anyone considering studies on body composition." Ecoscience

    " is a book that a researcher will want to check out and consult repeatedly in designing an ecological study...this is an extremely valuable book. There is no similar work that brings together multiple methods of non-destructive analysis for comparison and contrast...any library that supports an ecology community should definetly have this book on its shelves- or, more likely, in the hands of its patrons." E-Streams

  4. Han, J-D.J and SPEAKMAN, J.R. (2014)

    Targeting aging

    Journal of Genetics and Genomics

    Edited special issue on ageing

  5. SPEAKMAN, J.R. (2014)

    Pandas – dead end or dead wrong? and eleven other

    short stories from the frontiers of bioscience, 2014.

    Create space publishing

    ISBN-13: 978-1505548877
    ISBN-10: 150554887X

  6. SPEAKMAN, J.R. (2015)

    How dogs make us fall in love with them:

    and eleven other shrt stories from the

    frontiers of bioscience 2015.

    Createspace publishing

    ISBN-13: 978-1519681867

Unrefereed Review Papers and Unrefereed Invited Articles

  1. SPEAKMAN, J.R (1983)

    The searching behaviour of foraging redshanks Tringa totanus L.

    Wader Study Group Bulletin 37: 13-16

  2. SPEAKMAN, J.R. (1984)

    Why do redshank stop foraging when it is windy?

    Wader Study Group Bulletin 42:15

  3. Racey, P.A., SPEAKMAN, J.R. and swift, S.M. (1985)

    Bat studies in Scotland.

    Scottish Field Studies 1985: 31-35.

  4. Racey, P.A. and SPEAKMAN, J.R. (1985)

    Bats in Scotland

    Bat News (London) No. 3 Pages: 2p

  5. SPEAKMAN, J.R. and Smith, S.W.G. (1986)

    Meal ticket

    New Scientist 112: 73

  6. SPEAKMAN, J.R., Iason, GR & Grisley, MS (1986)

    Postdocs need a career structure

    Nature 326: 822

  7. SPEAKMAN, J.R. (1987)

    Mixed clutch of oystercatcher and lapwing eggs incubated by an oystercatcher.

    Scottish Birds 14: 184-185.

  8. SPEAKMAN, J.R. (1990)

    Bias in the collection of mussel shells opened by oystercatchers.

    Wader Study Group Bulletin 58: 48-49

  9. SPEAKMAN, J.R., Racey, P.A., McLean, J.A. and Entwistle, A.C. (1993)

    Six new records of Nathusius' pipistrelle Pipistrellus nathusii for Scotland.

    Scottish bats 2: 14-16

  10. SPEAKMAN, J.R., Entwistle, A.C.; McLean, J.A. (1993)

    Natterers bat (Myotis nattereri) in North east Scotland.

    Scottish Bats 2:11-12

  11. SPEAKMAN, J.R. (1993)

    Clustering in the emergence behavior of bats: Some pitfalls in analysis and how to overcome them

    Bat Research News 34: 49-54

  12. Cole, K.R and SPEAKMAN, J.R. (1993)

    Measurement of the resting metabolic rate of the Atlantic bottlenose dolphin (Tursiops truncatus).

    European Research on Cetaceans 7: 247-250

  13. SPEAKMAN, J.R., Racey, P.A.; Rydell, J. (1995)

    Second British record for the Northern Bat Eptesicus nilssonii: from a North Sea oil platform.

    Scottish Bats 3: 9

  14. Speakman and Roberts (intro to symposium) (1995)

    The doubly-labelled water method


  15. SPEAKMAN, J.R (2000)

    Comments on the proposed designation of neotypes for the nominal species Vespertilio pipistrellus Schreber, 1774 and V. pygmaeus Leach, 1825 (currently Pipistrellus pipistrellus and P. pygmaeus; Mammalia, Chiroptera).

    Bulletin of Zoological Nomenclature 57: 50 Published: 31 March 2000

  16. SPEAKMAN, J.R. and Selman, C. (2002)

    The free-radical theory of ageing. Invited article

    In “The Encyclopaedia of Ageing”

  17. Selman C, McLaren JS, Collins AR, SPEAKMAN, J.R. (2002)

    Voluntary exercise has only limited effects on activity of antioxidant enzymes and does not cause oxidative damage in a small mammal.

    Journal of Nutrition. 132: 1784S-1786S (Extended abstract)

  18. SPEAKMAN, J.R. (2003)

    Obesity. Part one. The biggest health threat facing mankind.

    The Biologist, 50: 11-14

  19. SPEAKMAN, J.R. (2003)

    Obesity. Part two. The physiology of body weight regulation.

    The Biologist, 50: 69-74

  20. SPEAKMAN, J.R. (2003)

    Obesity. Part three. Failed solutions and new ideas

    The Biologist, 50: 120-125

  21. SPEAKMAN, J.R. (2003)


    Article NG 0079 In “The Encyclopaedia of Energy” Editor in Chief C.J. Clevland.

    Elsevier press Los Angeles

  22. SPEAKMAN, J.R. (2003)

    Thermoregulation in vertebrates.

    Article A1824 in the Nature Encyclopaedia of Life Sciences

    Macmillan Press London

  23. SPEAKMAN, J.R. (2003)

    Thermoregulation in vertebrates : acclimation, acclimatisation and adaptation.

    Article A1825 in the Nature Encyclopaedia of Life Sciences

    Macmillan Press London

  24. Scantlebury, M., Clutton-Brock, T.H., and SPEAKMAN, J.R. (2005)

    Energetics of co-operative breeeding in meerkats.

    International Congress series 1275: 367-374.

  25. SPEAKMAN, J.R. (2005)

    Uncoupling mitochondria as a strategy for anti-ageing medicine

    Journal of European Anti-Ageing medicine and British Anti-Ageing Medical Journal 1: 27-29

    Invited contribution to launch issue

  26. Thomas, D. and SPEAKMAN, J.R. (2006)

    Physiological Ecology

    Section introduction pp 1-4 In Functional and evolutionary ecology of bats. Eds Zubaid, A., McCracken, G.F. and Kunz, T.H. Oxford University Press Oxford

  27. SPEAKMAN, J.R., Hambly, C., Mitchell, S., and Krol, E. (2007)

    Animal models of obesity.

    Obesity reviews 8: 55-61

  28. SPEAKMAN, J.R. (2006)
    The ice diet
    New Scientist 188: 65

  29. SPEAKMAN, J.R., Hambly, C., Mitchell, S.E. and Krol, E (2006)
    Animal models of obesity.
    DTI Foresight analysis. Tackling obesities

    (Invited report contribution)

  30. SPEAKMAN, J.R. and O’Rahilly, S. (2012)

    Fat - an evolving issue

    Disease models and mechanisms 5: 569-573

    Invited introductory article to special issue on evolution of obesity

  31. Roberts, S.B. and SPEAKMAN, J.R. (2013)

    Update on human caloric restriction research

    Advances in Nutrition. 4: 563-4.

    Invited summary of EB2013 session

  32. SPEAKMAN, J.R. (2013)

    Extra brainwashing at the weekend

    Newton (Dec 2013)

    Popular science article (in Chinese: translated by Lina Zhang)

  33. SPEAKMAN, J.R. (2014)

    Tool use: how low can it go?

    Newton (January 2014)

    Popular science article (in Chinese: translated by Lina Zhang)

  34. SPEAKMAN, J.R. (2014)

    Fears from the past

    Newton (February 2014)

    Popular science article (in Chinese: translated by Lina Zhang)

  35. SPEAKMAN, J.R. (2014)

    Should we abandon indirect calorimetry as a tool to diagnose energy expenditure? Not yet. Perhaps not ever.

    Molecular metabolism

    Invited commentary on paper by Burnett and Grobe (2014)

  36. SPEAKMAN, J.R. (2014)

    Cheap formations

    Newton (March 2014)

    Popular science article (in Chinese: translated by Lina Zhang)

  37. SPEAKMAN, J.R. (2014)

    Flashing too early

    Newton (April 2014)

    Popular science article (in Chinese: translated by Lina Zhang)

  38. SPEAKMAN, J.R. (2014)

    Lazy? It could be genetic and there could be a cure around the corner

    Newton (May 2014)

    Popular science article (in Chinese: translated by Lina Zhang)

  39. SPEAKMAN, J.R. (2014)

    Youthful blood

    Newton (June 2014)

    Popular science article (in Chinese: translated by Lina Zhang)

  40. SPEAKMAN, J.R. (2014)

    Crossing the boundary – Turing test

    Newton (July 2014)

    Popular science article (in Chinese: translated by Lina Zhang)

  41. SPEAKMAN, J.R. (2014)

    House work – it might save your life

    Newton (August 2014)

    Popular science article (in Chinese: translated by Lina Zhang)

  42. SPEAKMAN, J.R. (2014)


    In Earth Systems and Environmental Sciences

    Elsevier online encyclopedia.

  43. SPEAKMAN, J.R. (2014)

    Genes for running on thin air

    Newton (September 2014)

    Popular science article (in Chinese: translated by Lina Zhang)

  44. Han, J-D.J and SPEAKMAN, J.R. (2014)

    Targeting aging to decrease complex non-communicable human disease.

    Journal of Genetics and Genomics

    Invited introduction to special issue on aging

  45. SPEAKMAN, J.R. (2014)

    Stimulating memories

    Newton (October 2014)

    Popular science article (in Chinese: translated by Lina Zhang)

  46. SPEAKMAN, J.R. (2014)

    Cheetahs show how running costs more than walking

    Newton (November 2014)

    Popular science article (in Chinese: translated by Lina Zhang)

  47. SPEAKMAN, J.R. (2014)

    Do you need to be smart to do art?

    Newton (December 2014)

    Popular science article (in Chinese: translated by Lina Zhang)

  48. SPEAKMAN, J.R. (2015)

    Pandas – dead end or dead wrong

    Newton (January 2015)

    Popular science article (in Chinese: translated by Lina Zhang)

  49. SPEAKMAN, J.R. (2015)

    Beyond comprehension

    Newton (February 2015)

    Popular science article (in Chinese: translated by Lina Zhang)

  50. SPEAKMAN, J.R. (2015)

    The lottery you never want to win

    Newton (March 2015)

    Popular science article (in Chinese: translated by Lina Zhang)

  51. SPEAKMAN, J.R. (2015)

    When in Rome

    Newton (April 2015)

    Popular science article (in Chinese: translated by Lina Zhang)

  52. SPEAKMAN, J.R. (2015)

    The irresistible drug

    Newton (May 2015)

    Popular science article (in Chinese: translated by Lina Zhang)

  53. SPEAKMAN, J.R. (2015)

    Look into my eyes

    Newton (June 2015)

    Popular science article (in Chinese: translated by Lina Zhang)

  54. SPEAKMAN, J.R. (2015)

    Chocolate and weight loss

    Newton (July 2015)

    Popular science article (in Chinese: translated by Lina Zhang)

  55. SPEAKMAN, J.R. (2015)

    Pandas are cool.

    Newton (August 2015)

    Popular science article (in Chinese: translated by Lina Zhang)

  56. SPEAKMAN, J.R. (2015)

    Why are Panda’s so chilled? The clue is in the bamboo.

    The Conversation (July 2015)

    By July 2016 one year after publication this article had been read over 59,000 times.

  57. SPEAKMAN, J.R. (2015)

    Mathematically the most attractive

    Newton (August 2015)

    Popular science article (in Chinese: translated by Lina Zhang)

  58. SPEAKMAN, J.R. (2015)

    Who is the fattest of them all – and why?

    Newton (September 2015)

    Popular science article (in Chinese: translated by Lina Zhang)

  59. SPEAKMAN, J.R. (2015)

    Juice bar

    Newton (October 2015)

    Popular science article (in Chinese: translated by Lina Zhang)

  60. SPEAKMAN, J.R. (2015)

    No pain, no gain

    Newton (November, 2015)

    Popular science article (in Chinese: translated by Lina Zhang)

  61. SPEAKMAN, J.R. (2015)

    The Protein Compass

    Newton (December, 2015)

    Popular science article (in Chinese: translated by Lina Zhang)

  62. SPEAKMAN, J.R. (2016)

    Democracy – the cradle of mediocrity

    Newton (January 2016)

    Popular science article (in Chinese: translated by Lina Zhang)

  63. SPEAKMAN, J.R. (2016)

    A Life changing experience

    CAS Magazine

  64. SPEAKMAN, J.R. (2016)

    Keeping fit

    Newton (February 2016)

    Popular science article (in Chinese: translated by Lina Zhang)

  65. SPEAKMAN, J.R. (2016)

    The importance of being idle

    Newton (April 2016)

    Popular science article (in Chinese: translated by Lina Zhang)

  66. SPEAKMAN, J.R. (2016)

    Clearing out the junk

    Newton (March 2016)

    Popular science article (in Chinese: translated by Lina Zhang)

  67. SPEAKMAN, J.R. (2016)

    Shrinking birds

    Newton (May 2016)

    Popular science article (in Chinese: translated by Lina Zhang)

  68. SPEAKMAN, J.R. (2016)

    Women and Children first

    Newton (June 2016)

    Popular science article (in Chinese: translated by Lina Zhang)

  69. SPEAKMAN, J.R. (2016)

    Switching on and off the drive to eat

    Newton (July 2016)

    Popular Science article (in Chinese: translated by Lina Zhang)

  70. SPEAKMAN, J.R. Gorman, M.L., Mills, M.L.G. and Raath, J.P. (2016)

    Wild dogs and kleptoparasitism: some misunderstandings

    African Journal of ecology 54: 125-127

    Response to paper by

    Jongeling, T.B. and Koetsier, T (2014) The predicament of the African wild dog, Lycaon pictus, is less precarious than claimed. African Journal of Ecology 52: 466-470