Aging, Obesity, and Mortality: Misplaced Concern About Obese Older People?

Although there is widespread agreement that obesity (body mass index [BMI] ≥ 30 kg/m(2)) raises health risks, debate has ensued on whether obese older adults are also at greater risk. This study examines the effect of obesity on mortality for younger and older adults to determine whether the risk of obesity is lessened in later life. Data from a 20-year follow-up of a national sample of adults were used to examine the risk of obesity on mortality (N = 6,767). Cox models reveal that obesity raises mortality risk for adults of all ages, but this relationship is nearly twice as strong for persons younger than 50 years of age. Being slightly overweight in later life is associated with lower mortality risk, but obesity raises mortality risk, especially for ischemic heart disease. Obesity in middle age is a grave public health concern, but obesity in later life also merits attention.     

Maternal Obesity Is Associated with Alterations in the Gut Microbiome in Toddlers

Children born to obese mothers are at increased risk for obesity, but the mechanisms behind this association are not fully delineated. A novel possible pathway linking maternal and child weight is the transmission of obesogenic microbes from mother to child. The current study examined whether maternal obesity was associated with differences in the composition of the gut microbiome in children in early life. Fecal samples from children 18–27 months of age (n = 77) were analyzed by pyro-tag 16S sequencing. Significant effects of maternal obesity on the composition of the gut microbiome of offspring were observed among dyads of higher socioeconomic status (SES). In the higher SES group (n = 47), children of obese (BMI≥30) versus non-obese mothers clustered on a principle coordinate analysis (PCoA) and exhibited greater homogeneity in the composition of their gut microbiomes as well as greater alpha diversity as indicated by the Shannon Diversity Index, and measures of richness and evenness. Also in the higher SES group, children born to obese versus non-obese mothers had differences in abundances of Faecalibacterium spp., Eubacterium spp., Oscillibacter spp., and Blautia spp. Prior studies have linked some of these bacterial groups to differences in weight and diet. This study provides novel evidence that maternal obesity is associated with differences in the gut microbiome in children in early life, particularly among those of higher SES. Among obese adults, the relative contribution of genetic versus behavioral factors may differ based on SES. Consequently, the extent to which maternal obesity confers measureable changes to the gut microbiome of offspring may differ based on the etiology of maternal obesity. Continued research is needed to examine this question as well as the relevance of the observed differences in gut microbiome composition for weight trajectory over the life course.     

Life Years Lost Associated with Obesity-Related Diseases for U.S. Non-Smoking Adults

The objectives of this paper are to predict life years lost associated with obesity-related diseases (ORDs) for U.S. non-smoking adults, and to examine the relationship between those ORDs and mortality. Data from the National Health Interview Survey, 1997–2000, were used. We employed mixed proportional hazard models to estimate the association between those ORDs and mortality and used simulations to project life years lost associated with the ORDs. We found that obesity-attributable comorbidities are associated with large decreases in life years and increases in mortality rates. The life years lost associated with ORDs is more marked for younger adults than older adults, for blacks than whites, for males than females, and for the more obese than the less obese. Using U.S. non-smoking adults aged 40 to 49 years as an example to illustrate percentage of the life years lost associated with ORDs, we found that the mean life years lost associated with ORDs for U.S. non-smoking black males aged 40 to 49 years with a body mass index above 40 kg/m² was 5.43 years, which translates to a 7.5% reduction in total life years. White males of the same age range and same degree of obesity lost 5.23 life years on average – a 6.8% reduction in total life years, followed by black females (5.04 years, a 6.5% reduction in life years), and white females (4.7 years, a 5.8% reduction in life years). Overall, ORDs increased chances of dying and lessened life years by 0.2 to 11.7 years depending on gender, race, BMI classification, and age.     

Contributions of Incidence and Persistence to the Prevalence of Childhood Obesity during the Emerging Epidemic in Denmark

Background

Prevalence of obesity is the result of preceding incidence of newly developed obesity and persistence of obesity. We investigated whether increasing incidence and/or persistence during childhood drove the prevalence of childhood obesity during the emerging epidemic.

Methods

Height and weight were measured at ages 7 and 13 years in 192,992 Danish school children born 1930–1969. Trends in the incidence (proportion obese at 13 years among those not obese at 7 years) and persistence (proportion obese at 13 years among those obese at 7 years) across birth cohort periods (1930–41 with low stable prevalence of obesity, 1942–51 with increasing prevalence, 1952–69 with the higher, but stable prevalence) were investigated. Logistic regression was used to examine the associations between BMI at 7 years as a continuous trait, allowing interactions with the birth cohorts, and occurrence of obesity at 13 years.

Results

The prevalence of obesity was similar at 7 and 13 years and increased across birth cohorts in boys from around 0.1% to 0.5% and in girls from around 0.3% to 0.7%. The incidence of obesity between ages 7 and 13 years increased from 0.15% to 0.35% in boys and from 0.20% to 0.44% in girls. The persistence increased from 28.6% to 41.4% in boys and from 16.4% to 31.0% in girls. Despite a decrease over time, the remission of obesity occurred in >60% of obese children in the last birth cohort. However, the odds ratios of obesity at age 13 years in relation to the full range of BMI at 7 years remained unchanged across the birth cohort periods.

Conclusions/Significance

The development of the obesity epidemic in children was due to an increase in both incidence and persistence of obesity. Contrary to prevailing expectations, a large, although declining, proportion of children obese at an early age underwent remission during childhood.     

Obesity and early mortality in the United States

Objective:

Although obesity is a serious public health problem, there are few reliable measures of its health hazards in the United States. The objective of this study was to estimate how much earlier mortality is likely to occur for Americans who are obese (body mass index [BMI], ≥ 30).

Design and Methods:

Data from the National Health and Nutrition Examination Survey (NHANES) I (1971–1975), NHANES II (1976–1980), and NHANES III (1988–1994) for 37,632 participants who experienced 8,791 deaths during 15 years of follow‐up were prospectively analyzed. The relative risk of death from all causes and its advancement period, adjusted for covariates, were calculated. Stratification was used to investigate the effects of pre‐existing illness, smoking, and older age on the advancement period.

Results:

Compared to the participants of reference weight (BMI, 23 to <25 kg/m2), mortality was likely to occur 9.44 years (95% confidence interval [CI]: 0.72, 18.16) earlier for those who were obese (BMI, ≥ 30). For overweight (25 to <30 kg/m2), grade 1 obesity (BMI, 30 to <35) and grades 2–3 obesity (BMI, ≥ 35.0), the mortality was likely to occur earlier by 4.40 (?3.90, 12.70), 6.69 (?2.06, 15.43), and 14.16 (3.35, 24.97) years, respectively. These estimates apply to healthy nonsmoker young‐ and middle‐aged (21–55 years) adults, who constituted an estimated 32.8% of Americans with age of >21 years between 1988 and 1994. Without stratifying simultaneously for preexisting illness, smoking, and age, values of the advancement period for obesity were markedly smaller than those observed for healthy nonsmoker young and middle‐aged adults.

Conclusions:

For healthy nonsmokers young‐ and middle‐aged adults who constitute about one‐third of American adults, being obese is likely to hasten mortality by 9.44 years.

     

The effect of obesity on overall,circulatory disease- and diabetes-specific mortality

This paper explores the relationship between body mass and risk of death among US adults. The National Health Interview Survey-Multiple Cause of Death linked data set is used for the years 1987-1997, and Cox proportional hazard models are employed to estimate the association between obesity, as measured by the body mass index (BMI), and overall, circulatory disease-specific and diabetes-specific mortality. A U-shaped relationship is found between BMI and overall mortality. Compared with normal weight individuals, mortality during the follow-up period is 34% higher among obese class II individuals and 77% higher among obese class III individuals, controlling for age and sex. A J-shaped relationship exists between circulatory disease mortality and obesity, with a slightly higher risk of death for all categories of BMI. The relationship between BMI and diabetes mortality is striking. Compared with normal weight individuals, obese class I individuals are 2.8 times as likely to die, obese class II individuals are 4.7 times as likely to die, and obese class III individuals are 9.0 times as likely to die of diabetes during the follow-up period, controlling for age and sex. These results demonstrate that obesity heightens the risk of overall and circulatory disease mortality, and even more substantially increases the risk of diabetes mortality. These mortality findings, together with the substantial recent increases in obesity, lend urgency to public health programmes aimed at reducing the prevalence and consequences of obesity.     

Kin competition,natal dispersal and the moulding of senescence by natural selection

Most theoretical models for the evolution of senescence have assumed a very large, well mixed population. Here, we investigate how limited dispersal and kin competition might influence the evolution of ageing by deriving indicators of the force of selection, similar to Hamilton (Hamilton 1966 J. Theor. Biol. 12, 12–45). Our analytical model describes how the strength of selection on survival and fecundity changes with age in a patchy population, where adults are territorial and a fraction of juveniles disperse between territories. Both parent–offspring competition and sib competition then affect selection on age-specific life-history traits. Kin competition reduces the strength of selection on survival. Mutations increasing mortality in some age classes can even be favoured by selection, but only when fecundity deteriorates rapidly with age. Population structure arising from limited dispersal however selects for a broader distribution of reproduction over the lifetime, potentially slowing down reproductive senescence. The antagonistic effects of limited dispersal on age schedules of fecundity and mortality cast doubts on the generality of conditions allowing the evolution of ‘suicide genes’ that increase mortality rates without other direct pleiotropic effects. More generally, our model illustrates how limited dispersal and social interactions can indirectly produce patterns of antagonistic pleiotropy affecting vital rates at different ages.     

Diet composition and activity level of at risk and metabolically healthy obese american adults

Objective:

Obesity often clusters with other major cardiovascular disease risk factors, yet a subset of the obese appears to be protected from these risks. Two obesity phenotypes are described, (i) “metabolically healthy” obese, broadly defined as body mass index (BMI) ≥ 30 kg/m² and favorable levels of blood pressure, lipids, and glucose; and (ii) “at risk” obese, BMI ≥ 30 with unfavorable levels of these risk factors. More than 30% of obese American adults are metabolically healthy. Diet and activity determinants of obesity phenotypes are unclear. We hypothesized that metabolically healthy obese have more favorable behavioral factors, including less adverse diet composition and higher activity levels than at risk obese in the multi‐ethnic group of 775 obese American adults ages 40‐59 years from the International Population Study on Macro/Micronutrients and Blood Pressure (INTERMAP) cohort.

Design and Methods:

In gender‐stratified analyses, mean values for diet composition and activity behavior variables, adjusted for age, race, and education, were compared between metabolically healthy and at risk obese.

Results:

Nearly one in five (149/775 or 19%) of obese American INTERMAP participants were classified as metabolically healthy obese. Diet composition and most activity behaviors were similar between obesity phenotypes, although metabolically healthy obese women reported higher sleep duration than at risk obese women.

Conclusions:

These results do not support hypotheses that diet composition and/or physical activity account for the absence of cardiometabolic abnormalities in metabolically healthy obese.     

Life course pathways to adult-onset diabetes

Early life conditions, such as socioeconomic status (SES) and health, have the potential to set in motion multiple and reinforcing pathways that shape both the prevalence and onset of diabetes among older adults. Using data from the Health and Retirement Study (1998-2002) for persons age 51 years and older, we investigated the core mediating mechanisms linking early life conditions with diabetes prevalence in 1998 and onset over a 4-year follow-up period, focusing on adult achievement processes and obesity as key mechanisms. We found that father's education is negatively associated with diabetes prevalence for older men and women. However, no markers of early life SES are directly associated with older men's and women's onset of diabetes, and the negative effects of adult SES on diabetes onset pertain only to women. Early life health affects the onset of diabetes among women--but not the prevalence--and no evidence of this association was found for men. We found no evidence that obesity is an important mechanism connecting either early life or adult SES with diabetes development in men or women. We speculate that early life SES may accelerate the development of diabetes at younger ages, and that the pathways linking life course SES, early life health, and diabetes are partly gender-specific and biological in nature.     

Adult Obesity and the Burden of Disability throughout Life

Objective: To analyze the prevalence of disability throughout life and life expectancy free of disability, associated with obesity at ages 30 to 49 years. Research Methods and Procedures: We used 46 and 20 years of mortality follow‐up, respectively, for 3521 Original and 3013 Offspring Framingham Heart Study participants 30 to 49 years and classified as normal weight, overweight, or obese at baseline. Disability measures were available between 36 and 46 years of follow‐up for 1352 Original participants and at 20 years of follow‐up for 2268 Offspring participants. We measured the odds of disability in the Original cohort after 46 years follow‐up, and we estimated life expectancy with and without disability from age 50. Two disability measures were used, one representing limitations with mobility only and the second representing limitations with activities of daily living (ADL). Results: Obesity at ages 30 to 49 years was associated with a 2.01‐fold increase in the odds of ADL limitations 46 years later. Nonsmoking adults who were obese between 30 and 49 years lived 5.70 (95% confidence interval, 4.11 to 7.35) (men) and 5.02 (95% confidence interval, 3.36 to 6.61) (women) fewer years free of ADL limitations from age 50 than their normal‐weight counterparts. There was no significant difference in the total number of years lived with disability throughout life between those obese or normal weight, due to both higher disability prevalence and higher mortality in the obese population. Discussion: Obesity in adulthood is associated with an increased risk of disability throughout life and a reduction in the length of time spent free of disability, but no substantial change in the length of time spent with disability.     

Adaptive Content Biases in Learning about Animals across the Life Course

Prior work has demonstrated that young children in the US and the Ecuadorian Amazon preferentially remember information about the dangerousness of an animal over both its name and its diet. Here we explore if this bias is present among older children and adults in Fiji through the use of an experimental learning task. We find that a content bias favoring the preferential retention of danger and toxicity information continues to operate in older children, but that the magnitude of the bias diminishes with age and is absent in adults. We also find evidence that fitness costs likely impact the types of mistakes that participants make in their attributions of dangerousness and poisonousness. These results suggest that natural selection has shaped the way in which we learn and make inferences about unfamiliar animal species over ontogeny, and that future research is needed on how content biases may vary across the life course.     

Combined Impact of Cardiorespiratory Fitness and Visceral Adiposity on Metabolic Syndrome in Overweight and Obese Adults in Korea

Background

Obesity, especially visceral obesity, is known to be an important correlate for cardiovascular disease and increased mortality. On the other hand, high cardiorespiratory fitness is suggested to be an effective contributor for reducing this risk. This study was conducted to determine the combined impact of cardiorespiratory fitness and visceral adiposity, otherwise known as fitness and fatness, on metabolic syndrome in overweight and obese adults.

Methods

A total of 232 overweight and obese individuals were grouped into four subtypes according to their fitness level. This was measured by recovery heart rate from a step test in addition to visceral adiposity defined as the visceral adipose tissue area to subcutaneous adipose tissue area ratio (VAT/SAT ratio). Associations of fitness and visceral fatness were analyzed in comparison with the prevalence of metabolic syndrome.

Results

The high visceral fat and low fitness group had the highest prevalence of metabolic syndrome [Odds Ratio (OR) 5.02; 95% Confidence Interval (CI) 1.85–13.61] compared with the reference group, which was the low visceral adiposity and high fitness group, after adjustments for confounding factors. Viscerally lean but unfit subjects were associated with a higher prevalence of metabolic syndrome than more viscerally obese but fit subjects (OR 3.42; 95% CI 1.27–9.19, and OR 2.70; 95% CI 1.01–7.25, respectively).

Conclusions

Our study shows that visceral obesity and fitness levels are cumulatively associated with a higher prevalence of metabolic syndrome in healthy overweight and obese adults. This suggests that cardiorespiratory fitness is a significant modifier in the relation of visceral adiposity to adverse metabolic outcomes in overweight and obese individuals.     

Physical Inactivity Mediates the Association between the Perceived Exercising Behavior of Social Network Members and Obesity: A Cross-Sectional Study

Objectives

Social networks influence the spread of depression, health behaviors, and obesity. The social networks of older urban-dwelling adults were examined to assess whether physical inactivity mediated the association between social networks and obesity.

Methods

Data come from the Montreal Neighborhood Networks and Healthy Aging study (n = 2707). Self-reported height and weight were used to calculate body mass index (BMI) with obesity defined as a BMI≥30. A name generator/interpreter instrument was used to elicit participants’ core ties (i.e., alters), and assess whether alters exercised regularly and resided in participants’ neighborhoods. The International Physical Activity Questionnaire was used to measure physical inactivity. Separate multilevel logistic regression analyses were conducted for younger (18–54 years) and older (55 years plus) age groups to examine the association between the exercising behavior of alters and obesity. Ancillary analyses examined whether the residential location of alters was associated with obesity. Mediation analyses assessed whether physical inactivity mediated the association between alter exercising behavior and obesity. Models adjusted for participant socio-demographic and -economic characteristics.

Results

Among the older age stratum (55 years plus), physically inactive individuals were more likely obese (OR 2.14; 95% CIs: 1.48–3.10); participants who had more exercising alters were less likely obese (OR: 0.85; 95% CIs: 0.72–0.99). Physical inactivity mediated the association between exercising alters and obesity. Ancillary analyses showed that having exercising alters in the neighborhood compared to other locations tended to reduce the odds of obesity.

Conclusion

This work demonstrates the importance of social networks among older adults in facilitating a physically active lifestyle and reducing the odds of obesity. Such findings can inform the design of public health interventions that seek to improve the environmental conditions supporting the physical activity of older adults.     

Childhood Behavioral Problems Predict Young Adults' BMI and Obesity: Evidence From a Birth Cohort Stud

We examined whether behavioral problems in childhood and adolescence are associated with young adults' BMI and obesity, and tested whether childhood behavioral problems have a greater impact on young adults' obesity than adolescent behavioral problems. The data were from the Mater‐University of Queensland Study of Pregnancy (MUSP) and Its Outcomes, a population‐based birth cohort study commenced in Brisbane, Australia, in 1981. A subsample of 2,278 children for whom we had prospective information on their behavioral problems at ages 5 and 14 and measured BMI, and its categories (normal, overweight, and obese) at age 21 was chosen. Young adults who experienced behavioral problems at ages 5 or 14 had a greater average BMI and were more likely to be obese compared to young adults without behavioral problems at both ages. The childhood onset group was at greater risk of becoming obese by age 21 compared to the adolescent onset group (P = 0.04). These associations remained consistent after adjusting for a variety of potential covariates including maternal characteristics (i.e., demographics and life style), child dietary patterns, family meals, television (TV) watching, and participation in sports and exercise at 14 years. Childhood as well as persistent behavioral problems during childhood and adolescence predicts young adults' BMI and obesity. Although further studies are needed to confirm this association, there is a need for close monitoring of children presenting with behavioral problems.     

Disability prevalence among healthy weight,overweight, and obese adults

Objective:

Obesity is associated with adverse health outcomes in people with and without disabilities. However, little is known about disability prevalence among people who are obese. The purpose of this study is to determine the prevalence and type of disability among adults who are obese.

Design and Methods:

Pooled data from the 2003‐2009 National Health Interview Survey (NHIS) were analyzed to obtain national prevalence estimates of disability, disability type and obesity. The disability prevalence was stratified by body mass index (BMI): healthy weight (BMI 18.5‐<25.0), overweight (BMI 25.0‐<30.0), and obese (BMI ≥ 30.0).

Results:

In this pooled sample, among the 25.4% of US adults who were obese, 41.7% reported a disability. In contrast, 26.7% of those with a healthy weight and 28.5% of those who were overweight reported a disability. The most common disabilities among respondents with obesity were movement difficulty (32.5%) and work limitation (16.6%).

Conclusions:

This research contributes to the literature on obesity by including disability as a demographic in assessing the burden of obesity. Because of the high prevalence of disability among those who are obese, public health programs should consider the needs of those with disabilities when designing obesity prevention and treatment programs.     

Effect of current and midlife obesity status on mortality risk in the elderly

The primary purpose of this study was to determine whether current and midlife obesity status provide independent information on mortality risk in elderly persons. Analyses were based on 3,238 participants from the original Framingham Heart Study (FHS) cohort who lived to at least 70 years of age and who had BMI measures from when they were in their 50s. Within this group of 70-year olds, obesity based on current BMI was associated with a 21% increased risk of mortality (P = 0.019) whereas obesity in 70-year olds based on BMI measures obtained at around 50 years of age was associated with a 55% increased risk of mortality (P < 0.0001). Compared to 70-year olds who were nonobese at both 50 and 70 years of age, mortality risk was increased by 47% (P < 0.001) in those who were obese at both 50 and 70 years of age, increased by 56% (P < 0.001) in those who were obese at 50 years of age and nonobese at 70 years of age, and not significantly different (P > 0.9) in those who were nonobese at 50 years of age and obese at 70 years of age. In summary, in this cohort of elderly adults, midlife and current BMI had independent effects on mortality risk. Specifically, although mortality risk was increased in obese older adults who were already obese at midlife, this was not the case for newly obese older adults. Conversely, nonobese older adults who were obese at midlife had an increased mortality risk. These observations imply that it is imperative to consider an elderly adult's BMI in context of their BMI at midlife.     

Perceived Weight Discrimination and Obesity

Weight discrimination is prevalent in American society. Although associated consistently with psychological and economic outcomes, less is known about whether weight discrimination is associated with longitudinal changes in obesity. The objectives of this research are (1) to test whether weight discrimination is associated with risk of becoming obese (Body Mass Index≥30; BMI) by follow-up among those not obese at baseline, and (2) to test whether weight discrimination is associated with risk of remaining obese at follow-up among those already obese at baseline. Participants were drawn from the Health and Retirement Study, a nationally representative longitudinal survey of community-dwelling US residents. A total of 6,157 participants (58.6% female) completed the discrimination measure and had weight and height available from the 2006 and 2010 assessments. Participants who experienced weight discrimination were approximately 2.5 times more likely to become obese by follow-up (OR = 2.54, 95% CI = 1.58–4.08) and participants who were obese at baseline were three times more likely to remain obese at follow up (OR = 3.20, 95% CI = 2.06–4.97) than those who had not experienced such discrimination. These effects held when controlling for demographic factors (age, sex, ethnicity, education) and when baseline BMI was included as a covariate. These effects were also specific to weight discrimination; other forms of discrimination (e.g., sex, race) were unrelated to risk of obesity at follow-up. The present research demonstrates that, in addition to poorer mental health outcomes, weight discrimination has implications for obesity. Rather than motivating individuals to lose weight, weight discrimination increases risk for obesity.     

Meta‐analysis of the association between body mass index and health‐related quality of life among adults,assessed by the SF‐36

Objective:

Obesity is associated with impaired overall health‐related quality of life but individual studies suggest the relationship may differ for mental and physical quality of life. A systematic review using Medline, Embase, PsycINFO and ISI Web of Knowledge, and random effects meta‐analysis was undertaken.

Design and Methods:

Studies were included in the meta‐analysis if they were conducted on adults (defined as age >16 years), reported an overall physical and mental component score of the SF‐36, and, or both. Heterogeneity was assessed using I² statistics and publication and small study biases using funnel plots and Egger's test. Between‐study heterogeneity was explored using meta‐regression.

Results:

Eight eligible studies provided 42 estimates of effect size, based on 43,086 study participants. Adults with higher than normal body mass index had significantly reduced physical quality of life with a clear dose‐response relationship across all categories. Among class III obese adults, the score was reduced by 9.72 points (95% Confidence Interval 7.24, 12.20, P < 0.001). Mental quality of life was also significantly reduced among class III obese (?1.75, 95% confidence interval ?3.33, ?0.16, P = 0.031), but was not significantly different among obese (class I and class II) individuals, and was significantly increased among overweight adults (0.42, 95% confidence interval 0.17, 0.67, P = 0.001), compared to normal weight individuals. Heterogeneity was high in some categories, but there was no significant publication or small study bias.

Conclusions:

Different patterns were observed for physical and mental HRQoL, but both were impaired in obese individuals. This meta‐analysis provides further evidence on the impact of obesity on both aspects of health‐related quality of life.

     

Age-specific Effects of Novel Mutations in Drosophila Melanogaster II. Fecundity and Male Mating Ability

Evolutionary theories of senescence assume that mutations with age-specific effects exist, yet until now, there has been little experimental evidence to support this assumption. In this study, we allowed mutations to accumulate in an outbred, wild population of Drosophila melanogaster to test for age-specific differences in both male mating ability and fecundity. We assayed for age-specific effects of mutations after 10, 20, and 30 generations of mutation accumulation. For mating ability, we found the strongest effects of mutations in the first half of the life span after 20 generations, and at nearly all ages by generation 30. These results are qualitatively consistent with results from a companion study in which age-specific mortality was assayed on the same lines of D. melanogaster. By contrast, effects of fecundity were confined to late ages after 20 generations of mutation accumulation, but by generation 30, as with male mating ability, effects of novel mutations were distributed across all age classes. We discuss several possible explanations for the differences that we observe between generations within traits, and among traits, and the relevance for these patterns to models of aging as well as models of mate choice and sexual selection. This revised version was published online in July 2006 with corrections to the Cover Date.