500 q 2001 Blackwell Science Ltd Resveratrol, a natural dietary phytoalexin, possesses similar properties to hydroxyurea towards erythroid differentiation Christelle M. Rodrigue, 1 Nicole Arous, 1 Dora Bachir, 2 Juliette Smith-Ravin, 3 Paul-Henri Romeo, 1 Fre Âde Âric Galacteros 1,2 and Marie-Claude Garel 1 1 INSERM U474, Maternite  Port-Royal, ICGM Cochin, Paris, 2 Centre de la Dre Âpanocytose et des Thalasse Âmies, Ho Ãpital Henri Mondor AP±HP, Cre Âteil, France, and 3 Universite  des Antilles et de la Guyane, De Âpartement de Biologie Animale, Campus de Fouillole, Guadeloupe FWI) Received 7 August 2000; accepted for publication 8 January 2001 Summary. Resveratrol, a natural dietary polyphenol, has been postulated to be implicated in the cardioprotective effect of red wine and the low incidence of breast and prostate cancers among vegetarians and Orientals respec- tively. This compound inhibits ribonucleotide reductase as does hydroxyurea, the first therapeutic agent used in the treatment of sickle cell disease. Using the human erythro- leukaemic K562 cell line as an invitro model, we show here that 50 mmol/l of resveratrol induced a higher haemoglobin production sevenfold) in K562 cells than 500 mmol/l of hydroxyurea 3´5-fold). This erythroid differentiation was linked to a dose- and time-dependent inhibition of cell proliferation associated with an equivalent increased expression of p21 mRNA, but with a higher increased level of p21 protein sixfold) for cells treated with resveratrol than for those treated with hydroxyurea 1´5-fold). We also show that 50 mmol/l of resveratrol and 25 mmol/l of hydroxyurea induced variable but similar inhancements of fetal haemoglobin synthesis in cultured erythroid progeni- tors for the majority of the sickle cell patients studied. These inductions were linked to, but not correlated with, a variable decrease in erythroid burst-forming unit clone number. Taken together, these results show that resveratrol merits further investigations in sickle cell disease therapy. Keywords: K562 cells, erythroid differentiation, resveratrol, hydroxyurea, HbF. The 3, 5, 4 0 -trihydroxystilbene currently named resveratrol is a natural phytoalexin present in red wine and other constituents of the human diet Hain etal, 1990; Soleas etal, 1997). This polyphenol inhibits cyclooxygenase-1 activity and was implicated in chemopreventive action, which has been used to explain the low incidence of breast and prostate cancers among vegetarians and Orientals respec- tively Adlercreutz et al, 1995; Makela et al, 1995; Jang et al, 1997; Mgbonyebi et al, 1998; Carbo et al, 1999). Resveratrol is also a radical scavenger that prevents many heart disorders by inhibiting lipoprotein oxidation and platelet aggregation Frankel etal, 1993; Pace-Asciak etal, 1995; Jang etal, 1997; Rotondo etal, 1998). This molecule was associated with the low incidence of ischaemic heart disease observed in the French population, a phenomenon called the `French Paradox'. This was attributed to its presence in wine and to a moderate consumption of red wine Renaud & De Logeril, 1993; Soleas et al, 1997). Therefore, resveratrol seems to have a very broad range of biological properties Mizutani et al, 1998; Pendurthi et al, 1999), but little is known about the molecular basis of its activity. Recent studies have shown that resveratrol is a ribo- nucleotide reductase inhibitor Fontecave etal, 1998), much more effective than hydroxyurea HU), a molecule that has been used clinically against a variety of tumours. HU inhibits DNA synthesis and plays a role in cell growth inhibition associated with a reversible p53-independent induction of the cyclin-dependent kinase inhibitor CDKI) p21 Linke etal, 1996). HU is the first therapeutic agent for the treatment of sickle cell anaemia Charache etal, 1995), a haemoglobinopathy characterized by the polymerization of a mutated form of adult haemoglobin HbS) leading to erythrocyte sickling which plays an important part in the disease morbidity Bunn, 1997). Hydroxyurea has proven clinical benefit as it can increase fetal haemoglobin HbF) levels in the red cells of sickle cell patients inhibiting HbS polymer formation Platt etal, 1984; Charache etal, 1987; Steinberg etal, 1997; Maier-Redelsperger etal, 1998; Bunn, 1999). However, the increase in HbF is variable and some British Journal of Haematology , 2001, 113, 500±507 Correspondence: Dr Marie-Claude Garel, INSERM U474, Maternite  Port-Royal, 123 boulevard de Port-Royal, 75014 Paris, France. E-mail: garel@cochin.inserm.fr