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1. National Academy of Sciences. Nutrition during lactation. Washington, DC: National Academy Press, 1991.

2. Prentice AM. Can maternal dietary supplements help in preventing infant malnutrition? Acta Paediatr Scand 1991;S374:67-77.

3. Jensen RG, ed. Handbook of milk composition. New York: Academic Press, 1995.

4. Prentice AM, Prentice A. Evolutionary and environmental influences on human lactation. Proc Nutr Soc 1995;54:391 400.

5. Jensen RG. Lipids in human milk-composition and fatsoluble vitamins. In: Lebenthals E, ed. Textbook of gastroenterology and nutrition in infancy. New York: Raven Press, 1989:157-208.

6. Ballabriga A. Essential fatty acids and human tissue composition. An overview. Acta Paediatr 1994;S402: 63-8.

7. Carnielli VP, Luijendijk IHT, van Goudoever JB, Sulkers EJ, Boerlage AA, Degenhart HJ, Sauer PJJ. Feeding premature newborn infants palmitic acid in amounts and stereoisomeric position similar to that of human milk: effects on fat and mineral balance. Am J Clin Nutr 1995;61:103742.

8. Lonnerdal B. Biochemistry and physiological functions of human milk proteins. Am J Clin Nutr 1985;42:12991317.

9. Mata L. Breastfeeding and host defense. Front Gastrointest Res 1986;13:119-33.

10. Prentice A, Ewing G. Roberts SB, Lucas A, MacCarthy A, Jarjou LMA, Whitehead RG. The nutritional role of breastmilk IgA and lactoferrin. Acta Paediatr Scand 1987;76:592-8.

11. Mepham TB. Physiology of lactation. Milton Keynes, UK: Open University Press, 1987.

12. Kunz C, Rudloff S. Biological functions of oligosaccharides in human milk. Acta Paediatr S402;1994:903-12.

13. Koldovsky O. Hormonally active peptides in human milk. Acta Pediatr (Suppl) 1994;402:89-93.

14. Goldman AS, Goldblum RM. Defense agents in human milk. In: Jensens RG, ed. Handbook of milk composition. New York: Academic Press, 1995:727-45.

15. Prentice A, Prentice AM, Lamb WH. Mastitis in rural Gambian mothers and the protection of the breast by milk antimicrobial factors. Trans R Soc Trop Med Hyg 1985;79:90-5.

16. Wilde CJ, Prentice A, Peaker M. Breastfeeding: matching supply with demand in human lactation. Proc Nutr Soc 1995;54:401-6.

17. Schusdziarra V. Physiological role of beta-casomorphins. In: Picciano MF, Lonnerdals B. eds. Mechanisms regulating lactation and infant nutrient utilization. New York: WileyLiss, 1992:33748.

18. Stiehm RE. HIV transmission by breastfeeding: report of ten cases and review. In: Picciano MF, Lonnerdals B. eds. Mechanisms regulating lactation and infant nutrient utilization. New York: Wiley-Liss, 1992:179-88.

19. Astrup-Jensen A. Environmental and occupational chemicals. In: Bennett PN, ed. Drugs and human lactation. Amsterdam: Elsevier, 1988:551-73.

20. Chappell JE, Clandinin MT, Kearney-Volpe C. Trans fatty acids in human milk lipids: influence of maternal diet and weight loss. Am J Clin Nutr 1985;42:49-56.

21. Host A, Husby S. Osterballe O. A prospective study of cow's milk allergy in exclusively breastfed infants. Acta Paediatr Scand 1988;77:663-70.

22. Zarba A, Groopman JD. Biomarkers of aflatoxin exposure in humans: aflatoxin M1 in milk. In: Picciano MF, Lonnerdals B. eds. Mechanisms regulating lactation and infant nutrient utilization. New York: Wiley-Liss, 1992:451-5.

23. Berlin CM. Excretion of methylxanthines in human milk. Semin Perinatol 1981;5:389-94.

24. Dahlstrom A, Lundell B. Curvall M, Thapper L. Nicotine and cotinine concentrations in the nursing mother and her infant. Acta Paediatr Scand 1990;79:142-7.

25. DiLallo D, Bertollini R. CamposVenuti G. Risica S. Perucci CA, Simula S. Radioactivity in breast milk in Central Italy in the aftermath of Chernobyl. Acta Paediatr Scand 1987;76:530-1.

26. Bennett PN. Drugs and human lactation. Amsterdam: Elsevier, 1988.

27. Lazarus C, Edwards E. Radiopharmaceuticals. In: Bennett PN, ed. Drugs and human lactation. Amsterdam: Elsevier, 1988:495-549.

28. Cutting WAM. Breastfeeding and HIV infection: advice depends on the circumstances. Pediatr AIDS HIV Infect 1993;4:1-2.

29. Prentice AM, Prentice A. Energy costs of lactation. Annu Rev Nutr 1988;8:63-79.

30. George DE, DeFrancesca BA. Human milk in comparison to cow milk. In: Lebenthals E, ed. Textbook of gastroenterology and nutrition. New York: Raven Press, 1989:239-61.

31. Prentice A. Regional variations in the composition of human milk. In: Jensens RG, ed. Handbook of milk composition. New York: Academic Press, 1995:115-221.

32. Vorherr H. Human lactation and breast feeding. In: Larsons BL, ed. Lactation-a comprehensive treatise. New York: Academic Press, 1978:182-280.

33. Macy IG, Kelly HJ, Sloane RE. The composition of milks. Washington, DC: National Academy of Sciences, 1953.

34. Harzer G, Haug M, Bindels JG. Biochemistry of human milk in early lactation. Z Ernahrungswiss 1986;25:77-90.

35. Harzer G, Haschke F. Micronutrients in human milk. In: Renners E, ed. Micronutrients in milk and milkbased food products. London: Elsevier, 1989:125-238.

36. Michaelsen KF, Larsen PS, Thomsen BL, Samuelson G. The Copenhagen Cohort Study on Infant Nutrition and Growth: breast milk intake, human milk macronutrient content and influencing factors. Am J Clin Nutr 1994;59:600-11.

37. Prentice AM, Paul AA, Prentice A, Black AK, Cole TJ, Whitehead RG. Cross-cultural differences in lactational performance. In: Hamosh M, Goldmans AS, eds. Human lactation 2: Maternal and environmental factors. New York: Plenum Press, 1986:13-44.

38. Neville MC, Keller R. Seacat J. Lutes V, Neifert M, Casey C, Allen J. Archer P. Studies in human lactation: milk volumes in lactating women during the onset of lactation and full lactation. Am J Clin Nutr 1988; 48:1375-86.

39. Arthur PG, Smith M, Hartmann PE. Milk lactose, citrate and glucose as markers of lactogenesis in normal and diabetic women. J Pediatr Gastroenterol Nutr 1989;9:488-96.

40. Prentice A, Paul AA. Contribution of breast-milk to nutrition during prolonged breastfeeding. In: Atkinson SA, Hanson LA, Chandras RK, eds. Breastfeeding, nutrition, infection and infant growth in developed and emerging countries. St. John's, Newfoundland, Canada: ARTS Biomedical Publishers and Distributors, 1990:87-116.

41. Krebs NF, Reidinger CJ, Hartley S. Robertson AD, Hambidge KM. Zinc supplementation during lactation: effects on maternal status and milk zinc concentrations. Am J Clin Nutr 1995-,61:1030-6.

42. Prentice A, Prentice AM, Whitehead RG. Breast-milk fat concentrations of rural African women 1. Short-term variations within individuals. Br J Nutr 1981;45: 483-94.

43. Brown KH, Black RE, Robertson AD, Akhtar NA, Ahmed G. Becker S. Clinical and field studies of human lactation. Am J Clin Nutr 1982;35:745-54.

44. Laskey MA, Prentice A, Shaw J. Zachou T. Ceesay SM. Bresstmilk calcium concentrations during prolonged lactation in British and rural Gambian mothers. Acta Paediatr Scand 1990,79:507-12.

45. Prentice A, Prentice AM, Whitehead RG. Breast-milk fat concentrations of rural African women 2. Long-term variations within a community. Br J Nutr 1981; 45:495-503.

46. Prentice A. The effect of maternal parity on lactational performance in a rural African community. In: Hamosh M, Goldmans AS, eds. Human lactation 2: Maternal and environmental factors. New York: Plenum Press, 1986:165-73.

47. Prentice A, Jarjou LMA, Drury PJ, Dewit O. Crawford MA. Breast-milk fatty acids of rural Gambian mothers: effects of diet and maternal parity. J Pediatr Gastroenterol Nutr 1989;8:486-90.

48. Dewit O. Dibba B. Prentice A. Breastmilk amylase activity in English and Gambian mothers: effects of prolonged lactation, maternal parity and individual variations. Pediatr Res 1990;28:502-6.

49. Prentice A, Watkinson M, Prentice AM, Cole TJ, Whitehead RG. Breast-milk antimicrobial factors 11. Influence of season and prevalence of infection. Acta Paediatr Scand 1984;73:8039.

50. Bates CJ, Villard-Mackintosh L. Effects of low levels of riboflavin, vitamin C and vitamin A intake in Gambian lactating women. In: Picciano MF, Lonnerdals B. eds. Mechanisms regulating lactation and infant nutrient utilization. New York: Wiley-Liss, 1992:109-27.

51. Bates CJ, Prentice A. Breast milk as a source of vitamins, essential minerals and trace elements. Pharmacol Therapeut 1994;62:193-220.

52. Prentice AM, Goldberg GR, Prentice A. Body mass index and lactation performance. Eur J Clin Nutr 1994;48S:78-89.

53. Prentice A. Breastmilk calcium and phosphorus concentrations of mothers in rural Zaire. Eur J Clin Nutr 1991;45:611- 7,.

54. Salmenpera L, Perheentupa J. Nanto V, Siimes MA. Low zinc intake during exclusive breastfeeding does not impair growth. J Pediatr Gastroenterol Nutr 1995;18:361 -70.

55. Prentice A. Maternal calcium requirements during pregnancy and lactation. Am J Clin Nutr 1994;59S:47783.

56. Prentice A, Jarjou LMA, Cole TJ, Stirling DM, Dibba B. Fairweather-Tait S. Calcium requirements of lactating Gambian mothers: effects of a calcium supplement on breastmilk calcium concentration, maternal bone mineral content and urinary calcium excretion. Am J Clin Nutr 1995;65:58-67.

57. Prentice A, Dibba B. Jarjou LMA, Laskey MA, Paul AA. Is breastmilk calcium concentration influenced by calcium intake during pregnancy? Lancet 1994;344:4112.

58. Prentice A, Prentice AM, Cole TJ, Whitehead RG. Determinants of variations in breast-milk protective factor concentrations of rural Gambian mothers. Arch Dis Child 1983;58:518-22.

59. Department of Health. Weaning and the weaning diet. London: HMSO, 1994.

60. Prentice A. Breast feeding and the older infant. Acta Paediatr Scand 1991;S374:78-88.

61. Dagnelie PC, van Staveren WA, Verschuren SAJM, Hautvast JGAJ. Nutritional status of infants aged 4 to 18 months on macrobiotic diets and matched omnivorous control infants: a population-based mixed-longitudinal study. Eur J Clin Nutr 1989;43:311-23.


Discussion of paper by Prentice

Dr. Garza

Dr. Prentice, I also am intrigued by the calcium data. If you look at casein levels, are they comparable between the Gambia and Britain, or is the casein just not saturated? (I know that is not an appropriate term.)

Dr. Prentice

Clemens Kunz of the University of Dortmund in Germany measured the casein levels for us not long ago, and they seem to be very similar in American and Gambian mothers. In human milk, however, the main vehicle for calcium is citrate, not casein, and we are concentrating on the citrate levels first before going for the casein, as we are not expecting to see any major differences there.

Dr. McNeilly

Dr. Prentice, I have learned a lot. Could you tell me if nutrition might affect the rate of milk synthesis, or is that not shiftable?

Dr. Prentice

Dr. Hartmann could probably answer that question better than I. The only information we have is from our Gambian supplementation study, in which both unsupplemented and supplemented women produced the same amount of milk per day, but the supplemented women fed fewer times per day. So we think that it quite possibly has something to do with the mother's nutritional state, not necessarily with substrate supply affecting the storage capacity, as Dr. Hartmann said. Perhaps the unsupplemented women had smaller storage capacity and had to feed much more frequently during the day to produce the same volume. This is speculation, because we have not done any work on it. Have you any comment, Dr. Hartmann?

Dr. Hartmann

Yes, I think that when substrate supply is limited, the babies have to feed very frequently, regardless of the storage capacity, to keep the rate of synthesis at the maximum. Providing the mothers with more energy from the diet somewhat increases their ability to produce milk. Then the baby might not suckle as frequently to get the volume of milk that it requires, so I don't think it is inconsistent.

Dr. Rasmussen

I would like to contribute the results from a supplementation trial in Guatemala that have been presented at meetings but haven't yet been published. We selected women at the end of pregnancy and the beginning of lactation based on mid-calf circumference, not on body-mass index (BMI), because BMI has the properties that you mentioned. In that supplementation trial, we gave mothers either a low-energy or a high-energy supplement; we didn't have a no-treatment group, because we didn't consider it ethical, given the state of the population. The women in the high-energy group produced more milk, and-of particular importance to this meeting-they exclusively breastfed their babies for a longer period. We were not in the field long enough to ask if there was an effect on fertility in these women (in other words, on the length of their amenorrhoea), but I think there were clearly changes in the way the women breastfed as a result of changes in their milk volume. So I think there may be important effects of supplementation that we may not have seen in the trials that we have had today.

Dr. Prentice

I don't want to detract from that at all, but I still feel that we may be talking about two different things: the biological limits on lactation and the whole area of the mother's well-being-the social factors that affect the way she and the baby interact to increase milk volume. It seems from the work we have been doing that most women can produce reasonable volumes of milk, and if they don't, there are other factors that are probably affecting it other than the capacity of the breast in most instances. I wanted to ask you if you had any macronutrient content data from the breastmilk-that was my brief this morning. Does it affect the composition of the milk?

Dr. Rasmussen

The only component we have looked at is energy. There are compensatory changes in energy, such that women who produce more milk have less energy-dense milk The effects on infant growth are expected to be minimal. But our statistical power was not calculated on measuring infant growth.

Dr. Prentice

I think I would have predicted that. In our supplementation studies we found that if we gave women essentially a higherfat diet, there was more fat in the breastmilk, but lactose was reduced. This has been seen in quite a lot of other studies as well. Thus, the mother's breastmilk fat concentrations may well be responsive either to her own diet or to her adipose tissue stores. We have seen it, other people have seen it, but some other studies have not, and so it is still relatively controversial. But whatever happens, total milk energy seems to stay remarkably similar.

Dr. Hartmann

Could I add that we looked at some immunologic factors in the same material, and lactoferrin remained the same in both groups.

Dr. Garza

What was the content of fat in the Gambian diet?

Dr. Prentice

It's about 20% to 25%.

Dr. Garza

In studies done with Salvador Villalpando and Nancy Butte in Mexican women consuming diets of lower fat content, we found much lower levels of fat in human milk that were not fully compensated for by changes in lactose.

Dr. Prentice

That were nor compensated for?

Dr. Garza

That's right. We saw the same directional changes you did, but the reduction in fat was much greater than the increase in lactose, so that in populations with very low levels of fat (in the 10% to 12% range), the breastmilk fat content may be influenced dramatically.

Dr. Prentice

I certainly could believe that fat intakes might be limiting at that very low level. Did you find a difference in the shift between the manufactured fatty acids and the dietary fatty acids?

Dr. Garza

We didn't look at fat composition. Dr. Villalpando, has fat composition been examined since those earlier results were available?

Dr. Villalpando

We are looking at those data now. We see an increase in the medium-chain-length fatty acids, but the milk energy increase was not that impressive.

Dr. Prentice

Mike Crawford long ago suggested the same thing from Tanzanian studies, but there the total fat tended to stay the same but with larger amounts of medium-chain-length fatty acids.

Professor Howie

I apologize that this question is a little lateral to what you have been talking about, but I want to ask about the concept that breastfeeding mothers may not have to increase their own diet in proportion to the amount of milk they are giving the baby. In other words, if a mother is breastfeeding, the total energy cost to the community is less than if she were feeding herself and her baby by artificial means. That concept was discussed quite a bit a year or two ago, and I wonder if you have any comments on that and on its potential importance for nutrient provision.

Dr. Prentice

I think I would still say the same thing. There do appear to be women who are not able to increase their food intake, that is, their energy intake, and there do seem to be relatively simple compensations that can occur: a reduction in activity, for example. We are still not really sure that in lactation there is much in the way of changes in costs like basal metabolic rate so on, although there may well be in pregnancy, but fairly minor and adjustments may be satisfactory to enhance efficiency. These include accessing energy from fat stores; many, but not all, women in that situation will lose some of their fat stores while they are lactating. Certainly, in the Gambia some women put on weight during lactation at the time of year when food in good supply. But they are obviously eating enough is for both weight gain and milk production. So, yes, I think we would still be saying the same, that there are compensations for the energy needs of lactation.

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