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11.3 Brief comms AM NEW 5/3/04 9:45 am Page 3brief communicationsCOMMUNICATIONS ARISINGthe abundance of mural-precursor cells responders will never reject any non-zeroallowed efficient recruitment and invest- Animal behaviour offer). Once some responders are willing toment of mural cells to the engineered ves- reject low offers, self-interest guaranteesInequity aversion insels, thereby enhancing their stability. that proposers will raise their offers.The vascular permeability of the engi- Tw o kinds of variation from the standardcapuchins?neered vessels is higher than that of normal, ultimatum game suggest that humans,1quiescent vessels, but is in the lower range rosnan and de Waal have shown that unlike capuchins, would not reject in Bros-of permeability values induced by various nan and de Waal’s experimental context.capuchin monkeys are more likely toangiogenic molecules (see supplementary Breject a cucumber slice after seeing Two versions of a reduced-form ultimatum6information). As in normal microcircula- game have been comparedthat another capuchin has received a more : the first was ation, arteriolar and venular sides of the attractive grape. In interpreting this find- standard game, except that proposers hadengineered vessels are readily identified by ing, the authors make a link to work in only two choices, an equitable allocation orthe pattern of blood flow. Local administra- humans on ‘inequity aversion’ and suggest a highly inequitable ...

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the abundance of mural-precursor cells allowed efficient recruitment and invest-ment of mural cells to the engineered ves-sels, thereby enhancing their stability. The vascular permeability of the engi-neered vessels is higher than that of normal, quiescent vessels, but is in the lower range of permeability values induced by various angiogenic molecules (see supplementary information). As in normal microcircula-tion, arteriolar and venular sides of the engineered vessels are readily identified by the pattern of blood flow. Local administra-tion of a vasoconstrictor, endothelin-1, prompted constriction of the engineered arterial vessels. The few surviving arterioles in the HUVEC-alone constructs were sig-nificantly less contractile than those in the combined constructs (see supplementary information). These results indicate that the vessels that formed in the co-implanta-tion construct have a better functionality. Engineered blood vessels have often been found to be immature and unstable6. Genes that enhance the survival and/or pro-liferation of vascular cells –– endothelial cells and mural cells –– can be introduced to extend the lifespan of the engineered ves-sels5,7,8, but these may prove to be onco-genic. We have created long-lasting blood vessels without such genetic manipulation. In addition to realizing a crucial step in tissue engineering, our system provides a platform for testing thein vivofunctions of factors that control angiogenesis, vasculo-genesis and vessel maturation. The likely existence of endothelial and of smooth-muscle progenitor cells in adults9,10indi-cates that these cells might serve as a source of autologous cells for engineering blood vessels by using the approach described here. Naoto Koike, Dai Fukumura, Oliver Gralla, Patrick Au, Jeffrey S. Schechner*, Rakesh K. Jain Edwin L. Steele Laboratory, Department of Radiation Oncology, Massachusetts General Hospital, Harvard Medical School, Boston, Massachusetts 02114, USA e-mail:jain@steele.mgh.harvard.edu *Department of Dermatology, Yale University School of Medicine, New Haven, Connecticut 06520-8059, USA 1. Orlidge, A. & D’Amore, P. A.J. Cell Biol.105,1455–1462 (1987). 2. Hirschi, K. K., Rohovsky, S. A. & D’Amore, P. A.J. Cell Biol. 141,805–814 (1998). 3. Darland, D. C. & D’Amore, P. A.Angiogenesis4,11–20 (2001). 4. Jain, R. K., Munn, L. L. & Fukumura, D.Nature Rev. Cancer2, 266–276 (2002). 5. Schechner, J. S.et al. Proc. Natl Acad. Sci. USA97,9191–9196 (2000). 6. Jain, R. K.Nature Med.9,685–693 (2003). 7. Yang, J.et al. Nature Biotechnol.19,219–224 (2001). 8. McKee, J. A.et al. EMBO Rep.4,633–638 (2003). 9. Yamashita, J.et al. Nature408,92–96 (2000). 10. Simper, D., Stalboerger, P. G., Panetta, C. J., Wang, S. & Caplice, N. M.Circulation106,1199–1204 (2002). Competing financial interests:declared none. Supplementary information communicationaccompanies the onNature’s website.
brief communications
COMMUNICATIONS ARISINGresponders will never reject any non-zero Animal behaviouroffer). Once some responders are willing to reject low offers, self-interest guarantees Inequity aversion inthat proposers will raise their offers. capuchins?Two kinds of variation from the standard ultimatum game suggest that humans, r d de Waal1 capuchins, would not reject in Bros- unlikehave shown that Buced-fed ons rfaev ooisrwTamutmrnoa  mnlaunistoct a cucumber slci efaet resiegnmon eynkars moe l erlekiot yejer capuchi nan and de Waal’s experimental context. that another capuchin has received a more game have been compared6: the first was a attractive grape. In interpreting this find- standard game, except that proposers had ing, the authors make a link to work in only two choices, an equitable allocation or humans on ‘inequity aversion’ and suggest a highly inequitable one; and the second (the that capuchins, like humans, may reject ‘impunity game’) was identical, except that rewards because they are averse to unequal if the responder rejected his offer, the pro-pay-offs. Here I argue that this interpreta- poser s pay-off remained unchanged (but tion suffers from three problems: the the responder received zero). In the first results contradict the predictions of the version, players were willing to reject inequity-aversion model that Bosnan and inequitable allocations, whereas in the sec-de Waal cite2 version (which parallels the capuchin; experimental results indicate ond that humans do not behave like capuchins situation), subjects never rejected. in similar circumstances; and the available Similar evidence comes from multi-evidence does not suggest that inequity lateral ultimatum games in which one pro-aversion is cross-culturally universal3–5 faces multiple responders (see. poser I consider these points in turn. Brosnan U. Fischbacher, C. M. Fong & E. Fehr, and de Waal link their findings to work www.iew.unizh.ch/wp/iewwp133.pdf). In showing that a wide range of experimental this set-up, as long as one of the responders behaviour in humans can be understood by accepts the offer, the proposer gets his pay-introducing a preference for equity into the off. As before, the responder’s ability to standard self-interested utility function. affect the proposer’s pay-off by rejecting The effect of introducing this non-selfish is mitigated by the other responders who preference is to cause individuals — under might accept. As predicted by inequity aver-certain circumstances — to give up some of sion, responders decrease their willingness their pay-off in order to decrease the gains of to reject, and proposers drop their offers other individuals. accordingly. Therefore, although both of Applying the Fehr–Schmidt inequity- these experimental patterns are consistent aversion model2cited by Brosnan and de with inequity aversion, both also seem to Waal to the capuchin experimental situation be at odds with Brosnan and de Waal’s predicts that capuchins should always eat capuchin findings: that is, they show that the cucumber. It does not predict that humans will not reject unless this affects the inequity-averse individuals will reject the other’s pay-off. food reward, which is what the monkeys Brosnan and de Waal also suggest that did. Rejecting the cucumber increases, not inequity aversion is probably a human uni-decreases, inequality. Moreover, the grape- versal, and they cite work that uses the ulti-receiving capuchins sometimes reached matum game in 15 small-scale societies3–5. If through the cage and stole their partner’s responder behaviour (willingness to reject discarded cucumber, exacerbating the low offers) is taken as the most direct mea-inequality. sure of inequity aversion, then our results do Consistent with inequity aversion in not support the universal claim. Although humans, the results from experimental vari- five societies do show evidence consistent ations of the ultimatum game suggest that with inequity aversion, three others show humans would not reject a reward unless evidence of trivially little or no inequity that rejection reduced the take of the indi- aversion. The remainder have so few vidual who received more. In the ultimatum low offers that no substantial claims game, two players are allotted a sum of can be made. money to divide. The first player — the ‘pro-Joseph Henrich poser’ — must offer a portion of the sum toDepartment of Anthropology, Emory University, the second player — the ‘responder’ — whoAtlanta, Georgia 30322, USA must then decide whether to ‘accept’ ore-mail: jhenric@emory.edu ‘reject’ the offer. If the responder accepts, he S. & de Waal, F. B. M.1. Brosnan,Nature425,297–299 (2003). gets the amount of the offer and the propos-2. Fehr, E. & Schmidt, K. M.Q. J. Econ.114,817–868 (1999). er receives the remainder. If he rejects, both3. Henrich, J.et al.(eds)Foundations of Human Sociality: Economic Experiments and Ethnographic Evidence from 15 players get zero. The game is played once,Small-Scale Societies(Oxford University Press, New York, in and players never learn their partner’s iden-the press). 4. Henrich, J.Am. Econ. Rev.90,973–980 (2000). u in the will-itintgy.nIensse qofitrye sapvoernsdioerns  ctaon  reexjepclta lffers 5. Henrich, J.et al. Am. Econ. Rev.91,73–78 (2001). ow o6. Bolton, G. E. & Zwick, R.Games Econ. Behav.10, (in contrast, pure self-interest predicts that95–121 (1995).
NATURE VOL 428 11 MARCH 2004 www.nature.com/nature ©  2004Nature Publishing Group
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