Little is known about the wellbeing and mobility limitation of older disability retirees. Personal and environmental factors, such as time spent in working life, may either exacerbate or mitigate the onset of mobility limitation in general population. We aimed to study perceived midlife work ability as a determinant of self-reported mobility limitation in old age among municipal employees who transitioned into non-disability and disability retirement.
4329 participants of the Finnish Longitudinal Study of Municipal Employees (FLAME) had retired during January 1985 and July 2000. They had data on retirement, perceived work ability in 1985, and self-reported mobility limitation (non-disability retirement n = 2870, men 39%; and diagnose-specific disability retirement n = 1459, men 48%). Self-reported mobility was measured in 1985, 1992, 1997 and 2009. The latest score available was used to assess the number of mobility limitation. Work ability was measured by asking the respondents to evaluate their current work ability against their lifetime best in 1985. Incidence rate ratios (IRRs) and 95% confidence intervals (CIs) for work ability predicting mobility limitation in non-disability and diagnose-specific disability retirement groups were calculated using Poisson regression models.
The prevalence of mobility limitation for those who transitioned into non-disability retirement (Incidence Rate, IR = 0.45, 95% CI = 0.44-0.46) was lower compared to those who retired due to disability (IR = 0.65, CI = 0.63-0.66). A one-point increase in the work ability score decreased the risk for having one more mobility limitation among non-disability and all diagnose-specific retirement groups (musculoskeletal disease, cardiovascular disease, mental disorder, and other diseases).
Better midlife work ability may protect from old age mobility limitation among those who retire due to non-disability and disability. Promoting work ability in midlife may lead to more independent, active aging, regardless of type of retirement.
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To examine which individual and environmental factors correlate with unmet physical activity need in old age and predict development of unmet physical activity need (the feeling that one's level of physical activity is inadequate and thus distinct from the recommended amount of physical activity) over a 2-year follow-up.
Observational prospective cohort study and cross-sectional analyses.
Community and research center.
A total of 643 community-living ambulatory people aged 75 to 81 took part in face-to-face interviews and examinations at baseline and 314 at the 2-year follow-up.
Unmet physical activity need and its potential individual and environmental correlates were assessed at baseline. Development of unmet physical activity need was assessed over the 2-year follow-up period.
At baseline, all participants were able to walk at least 500 m outdoors, but 14% perceived unmet physical activity need. Unmet physical activity need was more prevalent in those with musculoskeletal diseases, depressive symptoms, and mobility limitations. Hills in the nearby environment, lack of resting places, and dangerous crossroads correlated with unmet physical activity need at baseline; the association was especially strong in those with walking difficulties. Significant baseline predictors for incident unmet physical activity need (15%) included fear of moving outdoors, hills in the nearby environment, and noisy traffic.
Unmet physical activity need is common in ambulatory community-living older people who have health and mobility problems and report negative environmental features in their neighborhood. Solutions to overcome barriers to physical activity need to be developed to promote equal opportunities for physical activity participation.
The aim was to study whether perceived environmental barriers to outdoor mobility affect changes in sense of autonomy in participation outdoors among community-dwelling older people over a two-year period.
Community-dwelling people aged 75-90 years (n = 848) in central Finland were interviewed on two occasions, face-to-face at baseline and over the telephone two years later. Perceived environmental barriers to outdoor mobility were assessed using a 15-item structured questionnaire, and the sum scores categorized into tertiles (0, 1 and 2 or more barriers). Autonomy in participation outdoors was assessed with the 'Impact on Participation and Autonomy' (IPA) questionnaire using the autonomy outdoors subscale (score range 0-20, higher scores indicating more restricted autonomy).
Scores for autonomy in participation outdoors were available for 848 participants at baseline (mean 6.2, SD = 3.8) and for 748 participants at the two-year follow-up (mean 6.7, SD = 3.9). At baseline, those reporting multiple environmental barriers had the most restricted autonomy, while those reporting no environmental barriers had the least restricted autonomy (p
The aim of this study was to investigate the association of noise sensitivity with self-reported hearing disability and hearing levels, with consideration of the role of self-reported history of noise exposure and use of hearing protectors. The study is based on the Finnish Twin Cohort. In 1988, a noise questionnaire was sent to 1005 twin pairs, 1495 individuals (688 men, 807 women) replied. The age range was 31-88 years. Information on some potential confounders was obtained from the questionnaire in 1981 for the same individuals. A subsample of thirty-eight elderly women with noise sensitivity response from 1988 had audiometry data from 2000 to 2001. Noise sensitivity was associated with self-reported hearing disability among all subjects [odds ratio (OR) 1.55, 95% confidence interval (CI) 1.14-2.12] and among women (OR 1.90, 95% CI 1.19-3.04), but no-more significantly among men (OR 1.31, 95% CI 0.86-1.98). The association was primarily seen among younger subjects (50 years or less). The difference between noise sensitive and non-noise sensitive elderly women in the average of thresholds at frequencies of 0.5-4 kHz in the better ear was not significant (Pr = 0.18). Noise sensitivity did not modify the association of hearing disability with the self-reported history of occupational noise exposure. Noise sensitivity was associated with the use of hearing protectors at work. The study shows the importance of recognizing the noise sensitive in noise effect studies, since sensitivity in annoyance has implications in most of the effect categories.
To profile participants based on reported outdoor physical activity barriers using a data-driven approach, describe the profiles and study their association with unmet physical activity need.
Cross-sectional analyses of 848 community-dwelling men and women aged 75-90 living in Central Finland in 2012. Barriers to outdoor physical activity and unmet physical activity need were enquired with a questionnaire. The latent profiles were identified by profiling participants into latent groups using a mixture modeling technique on the multivariate set of indicators of outdoor physical activity barriers. A path model was used to study the associations of the profiles with unmet physical activity need.
Five barrier profiles were identified. Profile A was characterized with minor barriers, profile B with weather barriers, profile C with health and weather barriers, profile D with barriers concerning insecurity, health and weather; and profile E with mobility and health barriers. The participants in the profiles differed in the proportion of individual and environmental barriers. The risk for unmet physical activity need was highest among people whose severe mobility difficulties restricted their outdoor physical activity.
Outdoor physical activity barriers reflect the imbalance in person-environment fit among older people, manifested as unmet physical activity need.
Environmental barriers are associated with disability-related outcomes in older people but little is known of the effect of environmental barriers on mortality. The aim of this study was to examine whether objectively measured barriers in the outdoor, entrance and indoor environments are associated with mortality among community-dwelling 80- to 89-year-old single-living people.
This longitudinal study is based on a sample of 397 people who were single-living in ordinary housing in Sweden. Participants were interviewed during 2002-2003, and 393 were followed up for mortality until May 15, 2012.Environmental barriers and functional limitations were assessed with the Housing Enabler instrument, which is intended for objective assessments of Person-Environment (P-E) fit problems in housing and the immediate outdoor environment. Mortality data were gathered from the public national register. Cox regression models were used for the analyses.
A total of 264 (67%) participants died during follow-up. Functional limitations increased mortality risk. Among the specific environmental barriers that generate the most P-E fit problems, lack of handrails in stairs at entrances was associated with the highest mortality risk (adjusted RR 1.55, 95% CI 1.14-2.10), whereas the total number of environmental barriers at entrances and outdoors was not associated with mortality. A higher number of environmental barriers indoors showed a slight protective effect against mortality even after adjustment for functional limitations (RR 0.98, 95% CI 0.96-1.00).
Specific environmental problems may increase mortality risk among very-old single-living people. However, the association may be confounded by individuals' health status which is difficult to fully control for. Further studies are called for.
The aim was to study objectively assessed walkability of the environment and participant perceived environmental facilitators for outdoor mobility as predictors of physical activity in older adults with and without physical limitations. 75-90-year-old adults living independently in Central Finland were interviewed (n = 839) and reassessed for self-reported physical activity one or two years later (n = 787). Lower-extremity physical limitations were defined as Short Physical Performance Battery score =9. Number of perceived environmental facilitators was calculated from a 16-item checklist. Walkability index (land use mix, street connectivity, population density) of the home environment was calculated from geographic information and categorized into tertiles. Accelerometer-based step counts were registered for one week (n = 174). Better walkability was associated with higher numbers of perceived environmental facilitators (p
Cites: Am J Phys Med Rehabil. 2014 Oct;93(10):876-8324800719
Cites: J Aging Res. 2012;2012:62575822162808
Cites: Scand J Med Sci Sports. 2015 Aug;25(4):e368-7326152855
Cites: Int J Health Geogr. 2014 Mar 04;13:724588848
Stress has damaging effects on individual's health. However, information about the long-term consequences of mental stress is scarce.
This 28-year prospective cohort study examined on the associations between midlife stress and old age disability among 2,994 Finnish municipal professionals aged 44-58 years at baseline. Self-reported stress symptoms were assessed at baseline in 1981 and 4 years later in 1985 and perceived disability in 2009. For the baseline data, principal component analysis was used for differentiation into stress symptom profiles. The regression coefficient estimates for self-care disability (activities of daily living) and instrumental activities of daily living disability were estimated using left-censored regression. The odds ratios for mobility limitation were estimated using logistic regression.
Four midlife stress profiles were identified: negative reactions to work and depressiveness, perceived decrease in cognition, sleep disturbances, and somatic symptoms. We saw a clear gradient of increasing disability severity in old age for increasing intensity of midlife stress symptoms. In comparison with the participants with no stress symptoms, the extensively adjusted left-censored and logistic regression models showed that in old age, disability scores were almost 2-4 units higher and risk for mobility limitation was 2-3 times higher for those with constant stress symptoms in midlife.
Among occupationally active 44- to 58-year-old men and women, perceived stress symptoms in midlife correlated with disability 28 years later. Stress symptoms may be the first signs of decompensation of individual functioning relative to environmental demands, which may later manifest in disabilities.
To examine the genetic and environmental sources of variation in self-rated health (SRH) in older female twins and to explore the roles of morbidity, functional limitation, and psychological well-being as mediators of genetic and environmental effects on SRH.
Cross-sectional analysis of twin data.
One hundred two monozygotic and 115 dizygotic female twin pairs aged 63 to 76.
SRH was categorized as good, average, or poor. Morbidity was described using a physician-assessed disease-severity scale together with information about the presence of diabetes mellitus and cancer. Maximal walking speed measured over 10 m was used to assess physical functional limitation; the Mini-Mental State Examination and the Center for Epidemiologic Studies Depression Scale were used to characterize psychological well-being. The contributions of genetic and environmental (defined as familial (shared by siblings) or nonshared (unique to each sibling)) effects were assessed using univariate and multivariate structural equation modeling of twin data.
SRH did not have its own specific genetic effect but shared a genetic component in common with the genetic components underlying liability to disease severity, maximal walking speed, and depressive symptoms. It accounted for 64% of the variation in SRH, with environmental effects accounting for the remaining variation.
The current results suggest that there are no specific genetic effects on SRH but rather that genetic influences on SRH are mediated through genetic influences affecting chronic diseases, functional limitation, and mood.
The purpose of the present study was to examine, first, whether hearing acuity predicts falls and whether the potential association is explained by postural balance and, second, to examine whether shared genetic or environmental effects underlie these associations.
Hearing was measured using a clinical audiometer as a part of the Finnish Twin Study on Aging in 103 monozygotic and 114 dizygotic female twin pairs aged 63-76 years. Postural balance was indicated as a center of pressure (COP) movement in semi-tandem stance, and participants filled in a fall-calendar daily for an average of 345 days after the baseline.
Mean hearing acuity (better ear hearing threshold level at 0.5-4 kHz) was 21 dB (standard deviation [SD] 12). Means of the COP velocity moment for the best to the poorest hearing quartiles increased linearly from 40.7 mm(2)/s (SD 24.4) to 52.8 mm(2)/s (SD 32.0) (p value for the trend = .003). Altogether 199 participants reported 437 falls. Age-adjusted incidence rate ratios (IRRs) for falls, with the best hearing quartile as a reference, were 1.2 (95% confidence interval [CI] = 0.4-3.8) in the second, 4.1 (95% CI = 1.1-15.6) in the third, and 3.4 (95% CI = 1.0-11.4) in the poorest hearing quartiles. Adjustment for COP velocity moment decreased IRRs markedly. Twin analyses showed that the association between hearing acuity and postural balance was not explained by genetic factors in common for these traits.
People with poor hearing acuity have a higher risk for falls, which is partially explained by their poorer postural control. Auditory information about environment may be important for safe mobility.
Cites: Am J Otolaryngol. 1999 Nov-Dec;20(6):371-810609481
Cites: J Gerontol A Biol Sci Med Sci. 2008 Feb;63(2):171-818314453
To study which individual characteristics and environmental factors correlate with fear of moving outdoors and whether fear of moving outdoors predicts development of mobility limitation.
Observational prospective cohort study and cross-sectional analyses.
Community and research center.
Seven hundred twenty-seven community-living people aged 75 to 81 were interviewed at baseline, of whom 314 took part in a 3.5-year follow-up.
Fear of moving outdoors and its potential individual and environmental correlates were assessed at baseline. Perceived difficulties in walking 0.5 km and 2 km were assessed twice a year over a 3.5-year period.
At baseline, 65% of the women and 29% of the men reported fear of moving outdoors. Poor socioeconomic status; musculoskeletal diseases; slow walking speed; and the presence of poor street conditions, hills in the nearby environment, and noisy traffic correlated with fear of moving outdoors. At the first 6-month follow-up, participants with fear of moving outdoors had more than four times the adjusted risk (odds ratio (OR)=4.6, 95% confidence interval (CI)=1.92-11.00) of developing difficulties in walking 0.5 km and a three times greater adjusted risk (OR=3.10, 95% CI=1.49-6.46) for developing difficulty in walking 2 km compared with those without fear. The difference in the prevalence of walking difficulties remained statistically significant over the 3.5-year follow-up (P=.02 and P=.009, respectively).
Fear of moving outdoors is common in older adults and increases the risk of developing self-reported difficulties in walking 0.5 km and 2 km. Knowledge about individual and environmental factors underlying fear of moving outdoors and finding ways to alleviate fear of moving outdoors are important for community planning and prevention of disability.
We examined the association between perceived environmental barriers to outdoor mobility and loneliness among community-dwelling older people. In addition, we studied whether walking difficulties and autonomy in participation outdoors affected this association.
Cross-sectional analyses of face-to-face home interview data with 848 people aged 75-90 years (mean age: 80.1 years; 62% women) gathered within the "Life-Space Mobility in Old Age" (LISPE) project. Self-reports of loneliness, environmental barriers to outdoor mobility, and difficulties in walking 2 km were obtained with structured questionnaires. Autonomy in participation outdoors was assessed with the "Impact on Participation and Autonomy" questionnaire.
Altogether, 28% of participants reported experiencing loneliness sometimes or often. These participants also reported more difficulties in walking 2 km, restricted autonomy in participation outdoors, and more environmental barriers to outdoor mobility than people not experiencing loneliness. Snowy and icy winter conditions (odds ratio: 1.59 [95% confidence interval: 1.15-2.20]), long distances to services (odds ratio: 1.57 [1.00-2.46]), and hills in the nearby environment (odds ratio: 1.49 [1.05-2.12]) significantly increased the odds for loneliness, even after adjustments for walking difficulties, autonomy in participation outdoors, perceived financial situation, living alone, and health. Path modeling revealed that environmental barriers increased loneliness either through direct association or indirectly through restricted autonomy in participation outdoors.
Prospective studies should investigate whether removing environmental barriers to outdoor mobility improves autonomy in participation outdoors and alleviates loneliness among older people.
Previous studies in young and middle-aged men and women have shown that resting electrocardiographic (ECG) variables are influenced by genetic factors. However, the extent to which resting ECG variables are influenced by genetic factors in older women is unknown. Thus, the aim of this study was to estimate the relative contribution of genetic and environmental influences to individual differences in resting ECG variables among older female twins without overt cardiac diseases.
Resting ECG recordings were obtained from 186 monozygotic and 203 dizygotic twin individuals, aged 63-76 years. Quantitative genetic modeling was used to decompose the phenotypic variance in each resting ECG variable into additive genetic, dominance genetic, shared environmental, and unique environmental influences.
The results showed that individual differences in the majority of the resting ECG variables were moderately to highly explained by additive genetic influences, ranging from 32% for T axis to 72% for TV(5). The results also suggested dominance genetic influences on QRS duration, TV(1), and Sokolow-Lyon voltage (36%, 53%, and 57%, respectively). Unique environmental influences were important for each resting ECG variable, whereas shared environmental influences were detected only for QT interval and QTc.
In older women without overt cardiac diseases, genetic influences explain a moderate to high proportion of individual differences in the majority of the resting ECG variables. Genetic influences are especially strong for T-wave amplitudes, left ventricular mass, and hypertrophy indices, whereas other variables, including heart rate, intervals, and axes, are more affected by environmental influences.
In older adults, mobility limitations often coexist with overweight or obesity, suggesting that similar factors may underlie both traits. This study examined the extent to which genetic and environmental influences explain the association between adiposity and mobility in older women. Body fat percentage (bioimpedance test), walking speed over 10 m, and distance walked in a 6-min test were evaluated in 92 monozygotic (MZ) and 104 dizygotic (DZ) pairs of twin sisters reared together, aged 63-76 years. Genetic and environmental influences on each trait were estimated using age-adjusted multivariate genetic modeling. The analyses showed that the means (and s.d.) for body fat percentage, walking speed, and walking endurance were 33.2+/-7.3%, 1.7+/-0.3 m/s and 529.7+/-75.4 m, respectively. The phenotypic correlation between adiposity and walking speed was -0.32 and between adiposity and endurance it was -0.33. Genetic influences explained 80% of the association between adiposity and speed, and 65% of adiposity and walking endurance. Cross-trait genetic influences accounted for 12% of the variability in adiposity, 56% in walking speed, and 34% in endurance. Trait-specific genetic influences were also detected for adiposity (54%) and walking endurance (13%), but not speed. In conclusion, among community-living older women, an inverse association was found between adiposity and mobility that was mostly due to the effect of shared genes. This result suggests that the identification of genetic variants for body fat metabolism may also provide understanding of the development of mobility limitations in older women.
To cope at their homes, community-dwelling older people surviving a hip fracture need a sufficient amount of functional ability and mobility. There is a lack of evidence on the best practices supporting recovery after hip fracture. The purpose of this article is to describe the design, intervention and demographic baseline results of a study investigating the effects of a rehabilitation program aiming to restore mobility and functional capacity among community-dwelling participants after hip fracture.
Population-based sample of over 60-year-old community-dwelling men and women operated for hip fracture (n = 81, mean age 79 years, 78% were women) participated in this study and were randomly allocated into control (Standard Care) and ProMo intervention groups on average 10 weeks post fracture and 6 weeks after discharged to home. Standard Care included written home exercise program with 5-7 exercises for lower limbs. Of all participants, 12 got a referral to physiotherapy. After discharged to home, only 50% adhered to Standard Care. None of the participants were followed-up for Standard Care or mobility recovery. ProMo-intervention included Standard Care and a year-long program including evaluation/modification of environmental hazards, guidance for safe walking, pain management, progressive home exercise program and physical activity counseling. Measurements included a comprehensive battery of laboratory tests and self-report on mobility limitation, disability, physical functional capacity and health as well as assessments for the key prerequisites for mobility, disability and functional capacity. All assessments were performed blinded at the research laboratory. No significant differences were observed between intervention and control groups in any of the demographic variables.
Ten weeks post hip fracture only half of the participants were compliant to Standard Care. No follow-up for Standard Care or mobility recovery occurred. There is a need for rehabilitation and follow-up for mobility recovery after hip fracture. However, the effectiveness of the ProMo program can only be assessed at the end of the study.
Current Controlled Trials ISRCTN53680197.
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The aim of this study was to analyze whether the associations between perceived environmental and individual characteristics and perceived walking limitations in older people differ between those with intact and those with poorer lower extremity performance.
Persons aged 75 to 90 ( N = 834) participated in interviews and performance tests in their homes. Standard questionnaires were used to obtain walking difficulties; environmental barriers to and, facilitators of, mobility; and perceived individual hindrances to outdoor mobility. Lower extremity performance was tested using Short Physical Performance Battery (SPPB).
Among those with poorer lower extremity performance, the likelihood for advanced walking limitations was, in particular, related to perceived poor safety in the environment, and among those with intact performance to perceived social issues, such as lack of company, as well as to long distances.
The environmental correlates of walking limitations seem to depend on the level of lower extremity performance.
This study examined the relative contribution of genetic and environmental effects on maximal leg extensor power and also investigated whether leg extensor power and maximum voluntary isometric knee extensor strength share a genetic component.
Muscle functions were measured as part of the Finnish Twin Study on Aging in 101 monozygotic (MZ) and 116 dizygotic (DZ) female twin pairs aged 63-76 yr. Leg extensor power was measured using the Nottingham Leg Extensor Power Rig and maximum voluntary isometric knee extensor strength using an adjustable dynamometer chair. The analyses were carried out using the maximum likelihood method in Mx-program on the raw data set.
A bivariate Cholesky decomposition model showed that leg extensor power and isometric knee extensor strength shared a genetic component in common, which accounted for 32% of the total variance in leg extensor power and 48% in isometric knee extensor strength. In addition, power and strength had a nonshared environmental effect in common accounting for four percent of the variance in power and 52% in strength. Remaining variance for leg extensor power was due to trait-specific shared and nonshared environmental effects.
Observed genetic effect in common for leg extensor power and maximum voluntary isometric knee extensor strength indicated that these two traits are regulated by the same genes. However, also environmental effects have a significant role in explaining the variability in power and strength.
The aim of the present study was to examine the contribution of genetic and environmental factors to depressive symptoms among older women. The participants were 102 monozygotic and 115 dizygotic female twin pairs aged 64 to 76 years. Depressive symptoms were assessed by the Center for the Epidemiologic Studies Depression Scale. The contribution of genetic and environmental effects was estimated for the constructed depressiveness factor and for the subscales which were depressed mood, psychomotor retardation, lack of wellbeing and interpersonal difficulties. Of the variance in depressiveness, shared environmental influences accounted for 39% and nonshared environmental influences 61%. For the subscales, 24% to 62% of the variance was explained by individual, and 13% to 23% by shared, environmental factors. Lack of wellbeing had its own moderate additive genetic effect explaining 30% of the variance. This study showed that in older women predominantly environmental factors underlay individual differences in depressiveness; however, the factors varied to some extent between dimensions measured by the subscales.
A crucial issue for the sustainability of societies is how to maintain health and functioning in older people. With increasing age, losses in vision, hearing, balance, mobility and cognitive capacity render older people particularly exposed to environmental barriers. A central building block of human functioning is walking. Walking difficulties may start to develop in midlife and become increasingly prevalent with age. Life-space mobility reflects actual mobility performance by taking into account the balance between older adults internal physiologic capacity and the external challenges they encounter in daily life. The aim of the Life-Space Mobility in Old Age (LISPE) project is to examine how home and neighborhood characteristics influence people's health, functioning, disability, quality of life and life-space mobility in the context of aging. In addition, examine whether a person's health and function influence life-space mobility.
This paper describes the study protocol of the LISPE project, which is a 2-year prospective cohort study of community-dwelling older people aged 75 to 90 (n?=?848). The data consists of a baseline survey including face-to-face interviews, objective observation of the home environment and a physical performance test in the participant's home. All the baseline participants will be interviewed over the phone one and two years after baseline to collect data on life-space mobility, disability and participation restriction. Additional home interviews and environmental evaluations will be conducted for those who relocate during the study period. Data on mortality and health service use will be collected from national registers. In a substudy on walking activity and life space, 358 participants kept a 7-day diary and, in addition, 176 participants also wore an accelerometer.
Our study, which includes extensive data collection with a large sample, provides a unique opportunity to study topics of importance for aging societies. A novel approach is employed which enables us to study the interactions of environmental features and individual characteristics underlying the life-space of older people. Potentially, the results of this study will contribute to improvements in strategies to postpone or prevent progression to disability and loss of independence.
Cites: Arch Phys Med Rehabil. 2001 Feb;82(2):210-611239312
The purpose of the present study was to examine the relative contribution of genetic and environmental effects on the air-conducted hearing threshold levels at low (0.125-0.5 kHz), mid (1-2 kHz), and high (4-8 kHz) frequencies separately for the better and worse hearing ear in older women. We also examined the distribution of audiogram configurations. Data was analysed using quantitative genetic modelling. As part of the Finnish twin study on aging (FITSA), hearing was measured in 103 monozygotic and 114 dizygotic female twin pairs aged 63-76 years. Approximately every third subject had a flat type, and two-thirds a descending type of audiogram configuration. No significant difference was observed in the distribution of audiogram configurations between zygosity groups. In the better ear, additive genetic effects accounted for 64%-74% of the total variance at different frequencies. For the worse ear, environmental effects were larger. Although overall heritability is rather constant across the frequency spectrum, it is noteworthy that at low and high frequencies frequency-specific genetic and environmental effects together accounted for the majority of the total variance.