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DOI: 10.1126/science.1198364 , 1447 (2011);331 Science

, et al.Gregory M. Walton Outcomes of Minority Students A Brief Social-Belonging Intervention Improves Academic and Health

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units (fig. S9) [paired t test, P < 0.02, n = 19 pairs; median distances from the granular layer were 0.45 mm (source units) and 1.05 mm (target units)].

The present study demonstrated that canoni- cal feed-forward signal flow across cortical layers during sensory coding reverse to the feed-back direction during memory retrieval phase, which sug- gests flexible recruitment of interlaminar connectivity depending on the cognitive demands in the monkey association cortices (Fig. 4C). We used CSD anal- ysis to estimate cortical layers (Fig. 1, C to E, and fig. S1), and the observed stimulus-evoked CSD profiles were quite similar to those in the primary sensory cortices (17, 27). For some penetrations, we observed that the current sink positioned super- ficially next to the earliest-sink contact exhibited larger peak amplitudes and much longer durations thanthatoftheearliestcurrentsink.Thisobservation might reflect the cytoarchitectural nature of A36 as a dysgranular cortex (28) as well as the direct inputs to the deepest part of the superficial layer, which is consistent with anatomical observations (29).

A recent study in the rat primary auditory cor- tex demonstrated that the direction of interlam- inar signal flow depends on the cortical “state”: Sensory-evoked responses were initiated in the thalamorecipient layers and then propagated to the superficial and deep layers, whereas in spontane- ously active “up-states,” neuronal activity was initiated in the deep layers and then propagated to the superficial layers (27). These state-dependent

changes in the interlaminar signal flows in rats are consistent with our results obtained in mon- keys performing a memory task. Together, these findings highlight the flexibility of cortical lam- inar circuits. Further experiments will be needed to determine whether such flexible interlaminar connectivity is also implemented and used in other cortical areas for other cognitive demands.

References and Notes 1. L. R. Squire, J. T. Wixted, R. E. Clark, Nat. Rev. Neurosci.

8, 872 (2007). 2. W. A. Suzuki, Neuron 61, 657 (2009). 3. E. A. Murray, T. J. Bussey, L. M. Saksida, Annu. Rev.

Neurosci. 30, 99 (2007). 4. Y. Miyashita, Science 306, 435 (2004). 5. Y. Naya, M. Yoshida, Y. Miyashita, J. Neurosci. 23, 2861

(2003). 6. N. K. Logothetis, D. L. Sheinberg, Annu. Rev. Neurosci.

19, 577 (1996). 7. G. Buzsáki, Neuron 68, 362 (2010). 8. S. H. Wang, R. G. Morris, Annu. Rev. Psychol. 61, 49, C1

(2010). 9. M. P. Witter, E. I. Moser, Trends Neurosci. 29, 671 (2006). 10. A. Bollimunta, Y. Chen, C. E. Schroeder, M. Ding, J. Neurosci.

28, 9976 (2008). 11. A. K. Engel, P. Fries, Curr. Opin. Neurobiol. 20, 156 (2010). 12. I. E. Ohiorhenuan et al., Nature 466, 617 (2010). 13. R. J. Douglas, K. A. Martin, Annu. Rev. Neurosci. 27, 419

(2004). 14. J. J. Nassi, E. M. Callaway, Nat. Rev. Neurosci. 10, 360

(2009). 15. R. C. Reid, J. M. Alonso, Curr. Opin. Neurobiol. 6, 475 (1996). 16. J. Csicsvari, B. Jamieson, K. D. Wise, G. Buzsáki, Neuron

37, 311 (2003). 17. C. E. Schroeder, P. Lakatos, Trends Neurosci. 32, 9 (2009).

18. A. K. Engel, W. Singer, Trends Cogn. Sci. 5, 16 (2001). 19. T. Hirabayashi, Y. Miyashita, J. Neurosci. 25, 10299 (2005). 20. M. Tomita, J. J. Eggermont, J. Neurophysiol. 93, 378 (2005). 21. C. A. Atencio, C. E. Schreiner, PLoS ONE 5, e9521 (2010). 22. T. Hirabayashi, D. Takeuchi, K. Tamura, Y. Miyashita,

J. Neurosci. 30, 10407 (2010). 23. J. M. Alonso, L. M. Martinez, Nat. Neurosci. 1, 395 (1998). 24. Materials and methods are available as supporting

material on Science Online. 25. S. Lefort, C. Tomm, J. C. Floyd Sarria, C. C. H. Petersen,

Neuron 61, 301 (2009). 26. N. Weiler, L. Wood, J. I. Yu, S. A. Solla, G. M. G.

Shepherd, Nat. Neurosci. 11, 360 (2008). 27. S. Sakata, K. D. Harris, Neuron 64, 404 (2009). 28. W. A. Suzuki, D. G. Amaral, J. Comp. Neurol. 463, 67 (2003). 29. K. S. Saleem, K. Tanaka, J. Neurosci. 16, 4757 (1996). 30. This work was supported by a Grant-in-Aid for Specially

Promoted Research from the Ministry of Education, Culture, Sports, Science and Technology (MEXT) to Y.M. (19002010) and Grant-in-Aid for Young Scientists from MEXT to T.H. (18700378); a grant from Takeda Science Foundation to Y.M.; and Japan Society for the Promotion of Science (JSPS) Research Fellowships for Young Scientists to D.T. (1811234) and K.T. (211438). This work was also supported in part by Global Center of Excellence Program from MEXT. We thank M. Takeda for discussions and comments and H. Kasahara, R. Fujimichi, T. Matsui, and K. W. Koyano for advice and help with experiments.

Supporting Online Material www.sciencemag.org/cgi/content/full/331/6023/1443/DC1 Materials and Methods SOM Text Figs. S1 to S9 Tables S1 and S2 References

3 November 2010; accepted 8 February 2011 10.1126/science.1199967

A Brief Social-Belonging Intervention Improves Academic and Health Outcomes of Minority Students Gregory M. Walton1* and Geoffrey L. Cohen1,2

A brief intervention aimed at buttressing college freshmen’s sense of social belonging in school was tested in a randomized controlled trial (N = 92), and its academic and health-related consequences over 3 years are reported. The intervention aimed to lessen psychological perceptions of threat on campus by framing social adversity as common and transient. It used subtle attitude-change strategies to lead participants to self-generate the intervention message. The intervention was expected to be particularly beneficial to African-American students (N = 49), a stereotyped and socially marginalized group in academics, and less so to European-American students (N = 43). Consistent with these expectations, over the 3-year observation period the intervention raised African Americans’ grade-point average (GPA) relative to multiple control groups and halved the minority achievement gap. This performance boost was mediated by the effect of the intervention on subjective construal: It prevented students from seeing adversity on campus as an indictment of their belonging. Additionally, the intervention improved African Americans’ self-reported health and well-being and reduced their reported number of doctor visits 3 years postintervention. Senior-year surveys indicated no awareness among participants of the intervention's impact. The results suggest that social belonging is a psychological lever where targeted intervention can have broad consequences that lessen inequalities in achievement and health.

A n important question facing society con- cerns the origins of inequalities between socially marginalized and nonmarginal-

ized groups. Among the most consequential of inequalities is the poorer school and health out- comes experienced by African Americans, Latino

Americans, and other non-Asian ethnic minor- ities relative to European Americans. These dif- ferences occur at all levels of socioeconomic status (1–3).

Although many structural factors contribute to these inequalities, the present research exam-

ines a psychological factor: concern about social belonging. Social belonging—a sense of having positive relationships with others—is a fundamen- tal human need (4, 5). Social isolation, loneliness, and low social status harm not only subjective well-being (6) but also intellectual achievement (7) and immune function and health (8–11). Even a single instance of exclusion can undermine well- being (12, 13), intelligence quotient (IQ) test per- formance, and self-control (14).

Members of socially stigmatized groups, such as African Americans, may be relatively more uncertain about their social belonging in main- stream institutions like school and work (7). Be- cause their ethnic group is often negatively stereotyped and marginalized, they may be un- sure of whether they will be fully included in positive social relationships in these settings (2). As the sociologist Erving Goffman wrote, “The central feature of the stigmatized individual’s sit- uation in life…is a question of…‘acceptance’” (15). Uncertainty about belonging, especially when chronic, can undermine minorities’ performance (7, 16) and health (3, 17, 18). Social belonging may thus constitute a psychological lever where targeted intervention could yield broad benefits.

1Department of Psychology, Stanford University, Stanford, CA 94305, USA. 2School of Education and Graduate School of Business, Stanford University, Stanford, CA 94305, USA.

*To whom correspondence should be addressed. E-mail: [email protected]

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Such an intervention is reported here. Critical to its rationale is the insight that it is people’s subjective interpretations of the quality of their relationships, more so than the objective number or attributes of those relationships, that strongly affects well-being (5, 19). The present interven- tion, delivered to students during the challenging transition to college, was designed to encourage nonthreatening interpretations of adversity. During the transition to a new school, students can face frequent social setbacks and feelings of isolation. Their well-being and performance may depend, to a great extent, on whether they construe such experiences as evidence that they do not belong.

Because African-American students experi- ence relatively greater uncertainty about their belonging in school, they were expected to ben- efit from the intervention more than nonminority students (7). Further if, as we intended, the in- tervention triggered an enduring perceptual change in the encoding of social experience, its effects might persist over time. Short-term effects might compound into long-term effects through a re- cursive virtuous cycle, in which early perform- ance gains assure students of their belonging in school, which in turn improves their performance, in a repeating feedback loop (20). Students who feel more assured of their belonging may also initiate more social interactions and form better relationships on campus, facilitating their social integration and further benefiting their well- being, performance, and health (21).

The intervention was delivered to two cohorts of African-American (N = 49) and European- American (N = 43) students in the second semes- ter of their first year at a selective college (22, 23). To assess psychological responses to adversity, we asked participants to complete daily surveys in the first week after the intervention. To assess their long-term sense of belonging, health, and well-being, we asked them to complete an end- of-college survey 3 years later (completion rate 78.26%) (23). At the end of this survey, partici- pants were asked to authorize the release of their complete college academic transcript (authoriza- tion rate 97.22%) (23).

Participating students were randomly as- signed to either the belonging-treatment condi- tion or a control condition. In cohort 1, participants were recruited through convenience sampling; in cohort 2, through random sampling (23). An additional campus-wide control group was ob- tained by collecting the anonymized official grade-point averages (GPAs) of all European Americans (N = 1362) and African Americans (N = 194) in the same class years as participants but who did not participate in the study (23). This group was included in secondary analyses of GPA.

The intervention provided students with a narrative that framed social adversity in school as shared and short-lived (24). This message en- couraged students to attribute adversity not to fixed deficits unique to themselves or their ethnic group but to common and transient aspects of the

college-adjustment process. Upon arrival in a research lab, participants read a report of the ostensible results of a survey of more senior students at their school. Most students, the report indicated, had worried about whether they be- longed in college during the difficult first year but grew confident in their belonging with time. The survey results were said to be consistent across ethnic and gender groups. For instance, one sur- vey respondent was quoted as saying, “Freshman year even though I met large numbers of people, I didn’t have a small group of close friends…I was pretty homesick, and I had to remind myself that making close friends takes time. Since then…I have met people some of whom are now just as close as my friends in high school were” (23). Concerns about belonging were thus represented as common at first, as temporary, and as due to the challenging nature of the college transition.

To encourage participants to internalize the message, several steps exploited the “saying- is-believing effect”—the tendency to endorse messages that one has freely advocated (25). Par- ticipants were asked to write an essay describing how their own experiences in college echoed the experiences summarized in the survey. They then turned their essay into a speech, which they delivered to a video camera. These materials, participants were told, would be shown to future students to help ease their transition to college. Beyond facilitating internalization, this procedure averted the potential stigma of receiving an intervention, because it encouraged participants to see themselves as benefactors and not as beneficiaries (24, 26). In the control condition, the procedure was the same but the survey ad- dressed topics unrelated to belonging (e.g., change in social-political attitudes) (23). The intervention lasted about 1 hour.

Few analyses were moderated by cohort (i.e., no more than would be expected by chance alone and none involving the primary outcomes of GPA, health, or well-being). Thus, data from the two cohorts were combined to increase statistical

power. First, analyses examined the trajectory of students’ official GPA over time. In contrast to all other groups, African Americans in the control group showed no improvement in GPA from the fall of their freshman year, the semester before the intervention, through their sophomore, junior, and senior years [linear trend F < 1]. By contrast, the GPAs of intervention-treated African Amer- icans rose over time [for linear trend, F(1,34) = 13.79, P= 0.0007; for time × condition, F(1,34) = 4.16, P= 0.049]. The GPAs of European-American students also rose over time [F(1,29) = 6.88, P = 0.014] with no difference by condition [F < 1]. As Fig. 1 shows, the intervention set African Americans on an upward trajectory such that the gap between them and their European Americans classmates closed over time. By students’ senior year, the gap was cut by 79% (23).

Multiple regression, with student gender controlled, tested the effect of student race and condition (randomized control versus social- belonging treatment) on change in GPA—mean postintervention GPA (sophomore through senior years) minus mean preintervention GPA (fall term, first year) (23). There was no condition effect on preintervention GPA for either racial group [t values < 1] (table S1) (23). However, a significant condition effect on change in GPA emerged for African Americans [B = 0.30, t(65) = 2.54, P = 0.014] with no effect for European Americans[t<1][race×conditionB=–0.43, t(65)= –2.41, P = 0.019]. Virtually identical results were obtained when preintervention GPA was used as a covariate in an analysis of postintervention GPA [treatment effect for African Americans, B = 0.24, t(64) = 2.65, P = 0.010; treatment effect for European Americans, t < 1; race × condition: B = –0.31, t(64) = –2.27, P = 0.027]. The intervention closed the minority gap in 3-year GPA (SD = 0.36) from 0.29 points in the control condition to 0.14 points in the treatment condition, a 52% reduction.

Adding the campus-wide sample further sup- ports treatment efficacy. An agreement with uni-

Fig. 1. Raw GPA by student race, experimen- tal condition, and aca- demic term. Means are noncumulative and were combined across cohorts. Ranges in sample sizes and standard errors for European Americans are N = 25 to 33 and SE = 0.08 to 0.14; for African Americans, N = 30 to 37 and SE = 0.09 to 0.12.

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versity officials precludes the reporting of raw or adjusted means in this sample. To honor this agreement but present the results graphically, we performed analyses on residual postintervention GPA with preintervention GPA and gender con- trolled. Multiple regression on change in GPA and on raw postintervention GPA with preinter- vention GPA included as a covariate yield vir- tually identical results (23). As shown in Fig. 2A, treated African Americans had higher residual GPA scores than did African Americans campus- wide [B = 0.28, t(1620) = 3.97, P = 0.00008] and African Americans in therandomized controlgroup [B = 0.24, t(1620) = 2.62, P = 0.009]. The latter two groups did not differ [t < 1]. (Fig. 2A) (23).

Illustrating its broad impact, the interven- tion tripled the percentage of African Americans earning postintervention GPAs in the top 25% of their class, as measured by both residual and raw postintervention GPA, and tended to reduce the percentage of African Americans performing in the bottom 25% of their class on both indices (Fig. 2, B and C) (23).

What accounts for these treatment effects? Daily surveys, collected the week after the in- tervention, suggest that the intervention buffered African Americans against adversity (23). Among untreated African Americans, feelings of be- longing in school rose and fell with the degree of adversity students reported having experienced earlier that day and the day before. As adversity rose, belonging fell (mean within-subjects R = –0.45, derived from the average of individual participants’ within-subjects correlations, after each was subjected to a Fisher r-to-z trans- formation) (23). For treated African Americans, this relationship was reduced to nil [mean within- subjects R = 0.01], a significant reduction [t(59) =

2.99, P = 0.004]. In summary, the intervention robbed adversity of its symbolic meaning for African Americans, untethering their sense of belonging from daily hardship (27). Like treated African Americans, European Americans showed little relationship between adversity and belonging [for both conditions, mean within-subjects R = –0.09; condition difference, t < 1] [race × con- dition: t(59) = –2.04, P = 0.046].

These results provide a window into the shift in African-American students’ psychology caused by the intervention. This shift benefited their long-term performance. African Americans whose belonging was more robust to daily adversity— whose sense of belonging was relatively inde- pendent of their day-to-day adversity—showed greater improvement in their 3-year postinter- vention GPA [R = 0.51, P = 0.001] (23). The effect of the intervention in protecting African- Americans students’sense of belonging from daily adversity mediated its effect on their GPA (23). The intervention thus planted a change in social perception that, it appears, accompanied students long after the intervention ended to affect their performance in college.

Three years after the intervention, at the end of their college tenure, participants completed a survey to assess long-term effects on psychology, well-being, and health. Also, to assess whether the intervention operated beneath conscious aware- ness, we asked participants whether they remem- bered the intervention from 3 years previously, whether they thought it had affected them, and whether they agreed with its message. On no outcome did European Americans differ by con- dition [t values < 1.35, P values > 0.18]. African Americans, however, showed consistent treat- ment effects. The race × condition interaction

was not always significant, indicating that the treatment effect was not always larger for African Americans than for European Americans. Degrees of freedom vary because some measures were completed only by participants in cohort 2 (23).

If the intervention lessened how much Afri- can Americans’ belonging fluctuated with adver- sity, and if it did so by lessening how much they viewed campus life through the lens of race, then intervention-treated African Americans should (i) report greater stability and less uncertainty about their belonging in school {less agreement with items like, “When something bad happens, I feel that maybe I don’t belong at [school name]”} (7) and (ii) exhibit less cognitive accessibility of negative racial stereotypes and self-doubt (23). They did [self-reported belonging uncertainty, t(36) = –2.01, P = 0.052; accessibility of negative racial stereotypes, t(66) = –2.01, P = 0.049; accessibility of self-doubt, t(64) = –2.64, P = 0.010] (Fig. 3) (23).

Given the importance of social belonging for reducing stress and improving immune function and physical health (5, 8–11, 19) and the rel- atively poorer health experienced by African Americans, even those high in socioeconomic status (3), we examined effects on health. We assessed self-reported health, an important pre- dictor of morbidity and mortality (28), using the five-item general health component of the Med- ical Outcomes Study Short-Form Health Survey (23). We also asked participants how many times they had visited the doctor in the previous 3 months (cohort 1) or 1 month (cohort 2). African Americans reported being healthier and visit- ing the doctor less frequently in the treatment condition than in the control condition [t(32) = 2.48, P = 0.019 and t(63) = –2.23, P = 0.030,

Fig. 2. Cumulative academic performance from sophomore through senior year. Data were combined across cohorts. (A) Resid- ual sophomore-through-senior- year GPAs adjusted for student gender and preintervention (fall term, first year) GPA. Error bars represent T1 SE. Means repre- sent the degree to which stu- dents performed better (positive values) or worse (negative val- ues) than expected after the in- tervention in GPA units based on their gender and preintervention performance. (B) Percentage of students in the top and bottom 25% of their college class in residual postintervention GPA (i.e., postintervention GPA ad- justed for student gender and preintervention GPA). (C) Percent- age of students in the top and bottom 25% of their college class in raw postintervention GPA. For analytic details, see (23). Sample sizes for European Americans areNcampus-wide control group = 1362and Nexperimental groups = 33; for African Americans, Ncampus-wide control group = 194 and Nexperimental groups = 37.

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respectively] (Fig. 4, A and B). Whereas 60% of untreated African Americans had seen a doctor recently, only 28% of treated African Americans had [c2(1, N = 38) = 3.98, P = 0.046]. The race gap in self-reported health was eliminated in the treatment condition; interestingly, there was no gap for doctor visits (23). Future research should examine whether these patterns generalize to physiological and physical indicators of health (9) to assess the robustness of the effect beyond self-report outcomes and to identify biological pathways (11).

As further evidence of improved well-being, African Americans also scored higher on the Subjective Happiness Scale (23) [t(35) = 2.61, P = 0.013] (Fig. 4C). The happiness gap with Euro- pean Americans higher than African Americans disappeared in the treatment condition (23). The finding of a lasting positive impact on subjective happiness is noteworthy in light of research showing that individual happiness is relatively stable (6).

Participants were unaware of the interven- tion’s effect, suggesting that its efficacy did not depend on conscious awareness. Most students reported that they remembered participating in the study 3 years earlier (79% did). But when asked to describe “the most memorable and important” information they had learned in the study, few recalled the key content of the survey they had read (8% did), and few reported that the study had had “any” effect on their college ex- perience (14% did) (table S3). There was no condition difference on any of these outcomes for African Americans [c2(1, N = 37 to 38) values < 1.40, P values > 0.20]; treated African Americans ascribed no more effect to the study than un- treated African Americans. However, indirect mea- sures of recall and beliefs did show treatment effects. When asked to “guess” the process of change described in the survey they had read, more treated than untreated African Americans wrote that it concerned how students’ social ex- periences in college improve over time (50% versus 20%) [c2(1, N = 38) = 3.79, P = 0.052]. Additionally, treated students endorsed this mes- sage. When asked to describe their own experi- ences,moretreatedthanuntreatedAfricanAmericans volunteered that their own social experiences in college had improved over time (50% versus 20%) [c2(1, N = 38) = 3.79, P= 0.052]. The subtle nature of this intervention, with its influence occurring outside conscious awareness (29), may contribute to its efficacy. In some cases, conscious aware- ness may undo the effects of an intervention (30). More overt interventions risk sending the stig- matizing message that the beneficiaries are seen as in need of help. They may also cause resist- ance and reactance and undermine the sense of accomplishment people take in their success (26).

This study provides an experimental, longi- tudinal demonstration that a brief intervention to buttress feelings of social belonging can have significant effects on a wide range of important outcomes. The social-belonging intervention im-

proved the academic performance, self-reported health, and well-being of ethnic minority students over 3 years. The results suggest that inequality between marginalized and nonmarginalized groups arises not only from structural factors but also from concern about social belonging.

This concern can be mitigated by using a psychological remedy. The intervention provided students a nonthreatening frame for interpreting the daily challenges of school. By encouraging students to adopt this message as their own, the intervention made this message stick psycholog- ically. Along with other recent research, this study

highlights how the impact of adversity depends on its perceived meaning—how it is subjective- ly construed (24–26, 31–33). Changing subjec- tive construal is a fruitful avenue for intervention because many events are ambiguous and ame- nable to multiple interpretations. Moreover, a change in construal can become self-reinforcing. Students who feel confident in their belonging may experience the social world in a way that re- inforces this feeling. They may initiate more rela- tionships and thus obtain more opportunities for belonging and growth. Brief interventions that shore up belonging can thus promote performance

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Fig. 4. Self-reported health and happiness 3 years postintervention. Error bars represent T1 SE. Data were combined across cohorts where measures were completed by both cohorts. (A) Self-assessed general health (cohort 2). (B) Reported doctor visits in the previous month (cohorts 1 and 2). (C) Subjective happiness (cohort 2). The y axes in (A) and (C) represent the full range of each scale. The y axis in (B) represents about 2.00 standard deviations. Sample sizes in cohort 2 only are NEuropean Americans = 20 and NAfrican Americans = 23. Sample sizes in cohorts 1 and 2 are NEuropean Americans = 31 and NAfrican Americans = 38.

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and well-being even long after their delivery (7, 20, 25, 31, 34).

Importantly, the effectiveness of social- psychological interventions depends on factors in the context. Such interventions are unlikely to be effective in contexts without opportunities for learning. Also, because the present intervention works by changing people’s subjective interpre- tation of ambiguous events, it may be ineffective in openly hostile environments. Lastly, whether this intervention would work among younger or less-select students, or students from other mar- ginalized groups, is an important question for future research (20, 31, 34). These qualifica- tions noted, the results underscore the impor- tance of social belonging and subjective construal in contributing to social inequality and show how this insight can inform our collective efforts to promote equality in performance, health, and well-being.

References and Notes 1. K. Magnuson, J. Waldfogel, Steady Gains and Stalled

Progress: Inequality and the Black-White Test Score Gap (Russell Sage Foundation, New York, 2008).

2. C. M. Steele, S. J. Spencer, J. Aronson, in Advances in Experimental Social Psychology, M. P. Zanna, Ed. (Academic Press, San Diego, CA, 2002), pp. 379–440.

3. D. R. Williams, Ann. N. Y. Acad. Sci. 896, 173 (1999). 4. R. F. Baumeister, M. R. Leary, Psychol. Bull. 117, 497 (1995). 5. J. T. Cacioppo, B. Patrick, Loneliness: Human Nature and

the Need for Social Connection (Norton, New York, 2008) 6. S. Lyubomirsky, K. M. Sheldon, D. Schkade, Rev. Gen.

Psychol. 9, 111 (2005).

7. G. M. Walton, G. L. Cohen, J. Pers. Soc. Psychol. 92, 82 (2007).

8. L. F. Berkman, S. L. Syme, Am. J. Epidemiol. 109, 186 (1979). 9. S. Cohen, D. Janicki-Deverts, Perspect. Psychol. Sci. 4,

375 (2009). 10. G. E. Miller, N. Rohleder, S. W. Cole, Psychosom. Med.

71, 57 (2009). 11. B. N. Uchino, J. Behav. Med. 29, 377 (2006). 12. N. I. Eisenberger, M. D. Lieberman, K. D. Williams,

Science 302, 290 (2003). 13. K. D. Williams, in Advances in Experimental Social

Psychology, M. P. Zanna, Ed. (Academic Press, San Diego, CA, 2009), pp. 279–314.

14. R. F. Baumeister, J. M. Twenge, C. K. Nuss, J. Pers. Soc. Psychol. 83, 817 (2002).

15. E. Goffman, Stigma: Notes on the Management of Spoiled Identity (Simon and Schuster, New York, 1963), p. 8.

16. R. Mendoza-Denton, G. Downey, V. J. Purdie, A. Davis, J. Pietrzak, J. Pers. Soc. Psychol. 83, 896 (2002).

17. J. Blascovich, S. J. Spencer, D. Quinn, C. Steele, Psychol. Sci. 12, 225 (2001).

18. S. W. Cole, M. E. Kemeny, S. E. Taylor, J. Pers. Soc. Psychol. 72, 320 (1997).

19. S. Cohen et al., Health Psychol. 27, 268 (2008). 20. G. L. Cohen, J. Garcia, V. Purdie-Vaughns, N. Apfel,

P. Brzustoski, Science 324, 400 (2009). 21. T. D. Wilson, Science 313, 1251 (2006). 22. A previous paper reports relatively short-term results of

this intervention from the first cohort of students (7). 23. Information on materials and methods is available as

supporting material on Science Online. 24. T. D. Wilson, M. Damiani, N. Shelton, in Improving

Academic Achievement: Impact of Psychological Factors on Education, J. Aronson, Ed. (Academic Press, Oxford, 2002), pp. 91–110.

25. J. Aronson, C. B. Fried, C. Good, J. Exp. Soc. Psychol. 38, 113 (2002).

26. L. Ross, R. Nisbett, The Person and the Situation (McGraw-Hill, New York, 1991).

27. J. Crocker, C. T. Wolfe, Psychol. Rev. 108, 593 (2001).

28. E. L. Idler, Y. Benyamini, J. Health Soc. Behav. 38, 21 (1997).

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30. D. K. Sherman et al., J. Pers. Soc. Psychol. 97, 745 (2009).

31. L. S. Blackwell, K. H. Trzesniewski, C. S. Dweck, Child Dev. 78, 246 (2007).

32. J. J. Gross, R. A. Thompson, in Handbook of Emotion Regulation, J. J. Gross, Ed. (Guilford, New York, 2007), pp. 3–24.

33. J. P. Jamieson, W. B. Mendes, E. Blackstock, T. Schmader, J. Exp. Soc. Psychol. 46, 208 (2010).

34. G. L. Cohen, J. Garcia, N. Apfel, A. Master, Science 313, 1307 (2006).

35. We thank participating students and helpful administrators at an anonymous university for their involvement. We also thank P. Carr, E. Chen, C. Dweck, C. Logel, G. Miller, D. Paunesku, D. Sherman, S. Spencer, C. Steele, and D. Yeager for input and L. Cai, K. Crockett, J. Darwall, L. Egan, A. Master, M. Silverio, and R. Titcombe for assistance. This research was supported by grants from the American Psychological Association, the Society for the Psychological Study of Social Issues, and the Spencer Foundation awarded to G.M.W. and by a grant from the Russell Sage Foundation awarded to G.L.C.

Supporting Online Material www.sciencemag.org/cgi/content/full/331/6023/1447/DC1 Materials and Methods Tables S1 to S3 References

28 September 2010; accepted 14 February 2011 10.1126/science.1198364

Direct Interaction of RNA Polymerase II and Mediator Required for Transcription in Vivo Julie Soutourina,1* Sandra Wydau,1† Yves Ambroise,2 Claire Boschiero,1 Michel Werner1*

Gene transcription is highly regulated. Altered transcription can lead to cancer or developmental diseases. Mediator, a multisubunit complex conserved among eukaryotes, is generally required for RNA polymerase II (Pol II) transcription. An interaction between the two complexes is known, but its molecular nature and physiological role are unclear. We identify a direct physical interaction between the Rpb3 Pol II subunit of Saccharomyces cerevisiae and the essential Mediator subunit, Med17. Furthermore, we demonstrate a functional element in the Mediator–Pol II interface that is important for genome-wide Pol II recruitment in vivo. Our findings suggest that a direct interaction between Mediator and Pol II is generally required for transcription of class II genes in eukaryotes.

M ediator is a large multisubunit complex conserved in all eukaryotes (1). It acts as a link between specific protein regu-

lators and the RNA polymerase II (Pol II) tran- scription machinery (2). Mediator is required at most Pol II–transcribed gene promoters for reg- ulated gene expression (3–5). In Saccharomyces cerevisiae, Mediator is composed of 25 subunits and is organized in four structural modules: the tail, middle, head, and Cdk8 modules (6). A di- rect Mediator–Pol II interaction is indicated by previous copurification, coimmunoprecipita- tion (CoIP) experiments (7–9) and by in vivo form- aldehyde cross-linking (10). A model of the Mediator–Pol II complex determined by electron

microscopy (EM) at 35 Å resolution suggests that several Pol II subunits (Rpb1, 2, 3, 6, and 11) might contact the middle or the head of Mediator (11). It was recently suggested that Rpb4 and Rpb7 could also be implicated in interactions with Mediator (12–14). However, the requirement of a direct interaction between Mediator and Pol II for transcription activation has not been demon- strated. Moreover, the identity of the Mediator subunits contacting Pol II is unknown because of the low resolution of the Mediator structure. As a consequence, the mechanism by which Mediator recruits Pol II is poorly understood.

To identify the subunit(s) of Mediator that directly contact Pol II and determine the role of these interactions in transcription regulation, we usedaninvivophoto–cross-linkingapproachbased on the incorporation by the cell-translation sys- tem of photo-activable analogs of methionine and leucine in proteins [see supporting online material (SOM) text and figs. S1 and S2] (15, 16).

Because EM results (11) suggested potential interactions of 16 Mediator subunits belonging to the head (Med6, 8, 11, 17, 18, 19, 20, 22) and middle (Med1, 4, 5, 7, 9, 10, 21, 31) modules with Rpb1, 2, 3, 6, or 11 Pol II subunits, we immu- noprecipitated hemagglutinin (HA)–tagged pro- teins after in vivo cross-linking. Among the 80 pairwise contacts that we tested, only Myc-tagged Rpb3 and HA-tagged Med17 cross-linked (Fig. 1). These results demonstrate that the Rpb3 Pol II

1Commissariat à l’Energie Atomique et aux Energies Al- ternatives (CEA), iBiTec-S, Service de Biologie Intégrative et Génétique Moléculaire, F-91191 Gif-sur-Yvette cedex, France. 2CEA, iBiTec-S, Service de Chimie Bio-organique et Marquage, F-91191 Gif-sur-Yvette cedex, France.

*To whom correspondence should be addressed. E-mail: [email protected] (J.S.); [email protected] (M.W.) †Present address: Université Paris Descartes, EA 4065, Faculté des Sciences Pharmaceutiques et Biologiques, 75006 Paris, France.

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