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Additive Genetic (additive + genetic)
Terms modified by Additive Genetic Selected AbstractsNormal and Abnormal Personality Traits: Evidence for Genetic and Environmental Relationships in the Minnesota Study of Twins Reared ApartJOURNAL OF PERSONALITY, Issue 5 2002Kristian E. Markon ABSTRACT Recent studies have demonstrated substantial correlations between normal and abnormal personality traits. Yet little is known about how these correlations are mediated genetically and environmentally: Do normal and abnormal personality traits stem from the same underlying genes and environments? We addressed this question using data from 128 monozygotic and dizygotic twin pairs in the Minnesota Study of Twins Reared Apart (MISTRA). Additive genetic and nonshared environmental correlations between scales of the Minnesota Multiphasic Personality Inventory (MMPI),an index of abnormal personality,and the Multidimensional Personality Questionnaire (MPQ),an index of normal personality,were estimated. Results indicated that phenotypic correlations between normal and abnormal personality were mediated by genetic as well as environmental factors, although the magnitude of genetic mediation tended to be larger overall. Moreover, the patterns of phenotypic, genetic, and environmental relationships among the scales were similar, suggesting that influences on normal and abnormal personality act through systems common to both. It is suggested that future research focus on the neurogenetic substrates of these shared systems and how dysfunction in these systems influences development of disordered personality. [source] Heritability of Obesity-related Phenotypes and Association with Adiponectin Gene Polymorphisms in the Chinese National Twin RegistryANNALS OF HUMAN GENETICS, Issue 2 2010Juan Lee Summary The purpose of this study was to estimate the heritability of obesity-related phenotypes and investigate the association of adiponectin gene polymorphisms +45T>G and +276G>T with these measures in Chinese twins. 1260 twin pairs were recruited from two cities through the Chinese National Twin Registry System from 2001 to 2005. Two SNPs at the adiponectin locus (+45T>G and +276G>T) were genotyped. Structural equation modeling (SEM) was used to estimate heritability and the best-fitting variance component model. The regular association among all twins was analysed with generalised estimating equations (GEE). Sib-transmission/disequilibrium test (TDT) within dizygotic (DZ) twin pairs discordant for their genotype was performed using SEM. Additive genetic, common and unique environmental (ACE) model-based heritability of body mass index (BMI) was 61%, while additive genetic and unique environmental (AE)-model-based heritability of waist circumference (WC) and waist-hip ratio (WHR) were 75% and 61%, respectively. There was no association of adiponectin gene +45T>G and +276G>T genotypes with obesity-related phenotypes in all twins or discordant DZ twins. Our twins data did not support that there was an association between adiponectin gene polymorphisms +45T>G and +276G>T and the obesity-related phenotypes. Further studies are required to better understand the role of adiponectin gene polymorphisms in obesity. [source] Additive genetic and other sources of variation in growth traits of juvenile black bream Acanthopagrus butcheriAQUACULTURE RESEARCH, Issue 7 2005Robert G Doupé Abstract We used a factorial mating design to estimate the contribution of additive genetic, non-additive genetic and maternal effects to variation in growth traits of black bream Acanthopagrus butcheri (Munro) at 75, 130 and 180 days of age in the hatchery. Maternal genetic and environmental effects were greatest at 75 days of age, accounting for 9.1% of total phenotypic variance in wet weight, 11.4% of variance in standard length and 8.8% of variance in total length. At later ages maternal effects were much reduced, explaining 0.8,3.7% of phenotypic variance in growth traits. Additive genetic effects were greatest at 130 days of age, when they accounted for 17.4% of total phenotypic variance in wet weight, 21.4% of variance in standard length and 18.7% of variance in total length. Additive genetic effects were negligible (<1%) at 75 days of age and 4.8,5.5% of total phenotypic variance in growth traits at 180 days of age. Non-additive genetic effects (which also included common environmental effects because of families being raised in the same tank) explained 5.8,7.3% of total phenotypic variance in growth traits at 75 days of age, but were much smaller at later ages. Variable stocking densities among tanks up to 75 days significantly affected all growth trait measurements below 180 days of age. [source] Modeling the genetic and environmental association between peer group deviance and cannabis use in male twinsADDICTION, Issue 3 2009Nathan A. Gillespie ABSTRACT Background Peer group deviance (PGD) is linked strongly to liability to drug use, including cannabis. Our aim was to model the genetic and environmental association, including direction of causation, between PGD and cannabis use (CU). Method Results were based on 1736 to 1765 adult males from the Mid-Atlantic Twin Registry with complete CU and PGD data measured retrospectively at three time-intervals between 15 and 25 years using a life-history calendar. Results At all ages, multivariate modeling showed that familial aggregation in PGD was explained by a combination of additive genetic and shared environmental effects. Moreover, the significant PGD,CU association was best explained by a CU,PGD causal model in which large portions of the additive genetic (50,78%) and shared environmental variance (25,73%) in PGD were explained by CU. Conclusions Until recently PGD was assumed to be an environmental, upstream risk factor for CU. Our data are not consistent with this hypothesis. Rather, they suggest that the liability to affiliate with deviant peers is explained more clearly by a combination of genetic and environmental factors that are indexed by CU which sits as a ,risk indicator' in the causal pathway between genetic and environmental risks and the expression of PGD. This is consistent with a process of social selection by which the genetic and environmental risks in CU largely drive the propensity to affiliate with deviant peers. [source] EVOLUTIONARY POTENTIAL OF A LARGE MARINE VERTEBRATE: QUANTITATIVE GENETIC PARAMETERS IN A WILD POPULATIONEVOLUTION, Issue 4 2009Joseph D. DiBattista Estimating quantitative genetic parameters ideally takes place in natural populations, but relatively few studies have overcome the inherent logistical difficulties. For this reason, no estimates currently exist for the genetic basis of life-history traits in natural populations of large marine vertebrates. And yet such estimates are likely to be important given the exposure of this taxon to changing selection pressures, and the relevance of life-history traits to population productivity. We report such estimates from a long-term (1995,2007) study of lemon sharks (Negaprion brevirostris) conducted at Bimini, Bahamas. We obtained these estimates by genetically reconstructing a population pedigree (117 dams, 487 sires, and 1351 offspring) and then using an "animal model" approach to estimate quantitative genetic parameters. We find significant additive genetic (co)variance, and hence moderate heritability, for juvenile length and mass. We also find substantial maternal effects for these traits at age-0, but not age-1, confirming that genotype,phenotype interactions between mother and offspring are strongest at birth; although these effects could not be parsed into their genetic and nongenetic components. Our results suggest that human-imposed selection pressures (e.g., size-selective harvesting) might impose noteworthy evolutionary change even in large marine vertebrates. We therefore use our findings to explain how maternal effects may sometimes promote maladaptive juvenile traits, and how lemon sharks at different nursery sites may show "constrained local adaptation." We also show how single-generation pedigrees, and even simple marker-based regression methods, can provide accurate estimates of quantitative genetic parameters in at least some natural systems. [source] An examination of the overlap between genetic and environmental risk factors for intentional weight loss and overeatingINTERNATIONAL JOURNAL OF EATING DISORDERS, Issue 6 2009Tracey D. Wade PhD Abstract Objective: To further our understanding of how intentional weight loss (IWL) and overeating are related, we examined the shared genetic and environmental variance between lifetime IWL and overeating. Method: Interview data were available for 1,976 female twins (both members of 439 and 264 pairs of monozygotic and dizygotic twins, respectively), mean age = 40.61, SD = 4.72. We used lifetime diagnostic data for eating disorders obtained from a semistructured psychiatric telephone interview, examined in a bivariate twin analysis. Both lifetime behaviors were measured on a 3-point scale, where absence of IWL or overeating formed one anchor on the scale and lifetime anorexia nervosa (AN) and bulimia nervosa (BN) formed the opposite anchors, respectively. Results: In line with previous findings, a higher body mass index was significantly associated with the lifetime presence of IWL and/or overeating (odds ratio = 1.13, 95% confidence interval (CI): 1.08,1.19). The best fitting twin model contained additive genetic and nonshared environmental influence influencing both IWL and overeating, with correlations between these influences of 0.61 (95% CI: 0.35,0.92) and 0.24 (95% CI: 0.07,0.42), respectively. Discussion: About 37% of genetic risk factors were considered to overlap between IWL and overeating, and with only 6% of overlap between environmental risk factors. Thus, considerable independence of risk factors was indicated. © 2009 by Wiley Periodicals, Inc. Int J Eat Disord 2009 [source] Estimates of genetic parameters for conformation measures and scores in Finnhorse and Standardbred foalsJOURNAL OF ANIMAL BREEDING AND GENETICS, Issue 5 2010E. Schroderus Summary The aim of this study was to estimate genetic parameters for conformation measures and scores in the Finnhorse and the Standardbred foals presented in foal shows. Studied traits included height at withers and at croup, six subjectively evaluated conformation traits and overall grade. Data were from 10-year period (1995,2004) and consisted of 5821 Finnhorse foals (1,3 years old) with 7644 records and 2570 Standardbred foals (1,2 years old) with 2864 records. Variance components were estimated with REML , animal model using VCE4 program. The model included age class, year of judging, sex and region as fixed effects, and additive genetic, permanent environmental and residual as random effects. Estimates of heritability for measured traits were very high in both breeds (0.88,0.90). Estimates of heritability for conformation traits varied from 0.13 to 0.32 in the Finnhorse and from 0.06 to 0.47 in the Standardbred. In both breeds, estimates of heritability were lowest for hooves and movements at walk, and highest for type and body conformation among scored traits. Estimate of heritability for overall grade was in the Finnhorse 0.32 and in the Standardbred 0.34. Genetic correlations between overall grade and different conformation traits were 0.35,0.84 in the Finnhorse and 0.31,0.88 in the Standardbred. Thus, selection based on the overall grade would improve all studied characteristics. [source] Selection, structure and the heritability of behaviourJOURNAL OF EVOLUTIONARY BIOLOGY, Issue 2 2002D. G. Stirling Characters which are closely linked to fitness often have low heritabilities (VA/VP). Low heritabilities could be because of low additive genetic variation (VA), that had been depleted by directional selection. Alternatively, low heritabilities may be caused by large residual variation (VR=VP , VA) compounded at a disproportionately higher rate than VA across integrated characters. Both hypotheses assume that each component of quantitative variation has an independent effect on heritability. However, VA and VR may also covary, in which case differences in heritability cannot be fully explained by the independent effects of elimination-selection or compounded residual variation. We compared the central tendency of published behavioural heritabilities (mean=0.31, median=0.23) with morphological and life history data collected by Mousseau & Roff (1987). Average behavioural heritability was not significantly different from average life history heritability, but both were smaller than average morphological heritability. We cross-classified behavioural traits to test whether variation in heritability was related to selection (dominance, domestic/wild) or variance compounding (integration level). There was a significant three-way interaction between indices of selection and variance compounding, related to the absence of either effect at the highest integration level. At lower integration levels, high dominance variance indicated effects of selection. It was also indicated by the low CVA of domestic species. At the same time CVR increased disproportionately faster than CVA across integration levels, demonstrating variance compounding. However, neither CVR nor CVA had a predominant effect on heritability. The partial regression coefficients of CVR and CVA on heritability were similar and a path analysis indicated that their (positive) correlation was also necessary to explain variation in heritability. These results suggest that relationships between additive genetic and residual components of quantitative genetic variation can constrain their independent direct effects on behavioural heritability. [source] Evidence for an Interaction Between Age at First Drink and Genetic Influences on DSM-IV Alcohol Dependence SymptomsALCOHOLISM, Issue 12 2009Arpana Agrawal Background:, Research suggests that individuals who start drinking at an early age are more likely to subsequently develop alcohol dependence. Twin studies have demonstrated that the liability to age at first drink and to alcohol dependence are influenced by common genetic and environmental factors, however, age at first drink may also environmentally mediate increased risk for alcohol dependence. In this study, we examine whether age at first drink moderates genetic and environmental influences, via gene × environment interactions, on DSM-IV alcohol dependence symptoms. Methods:, Using data on 6,257 adult monozygotic and dizygotic male and female twins from Australia, we examined the extent to which age at first drink (i) increased mean alcohol dependence symptoms and (ii) whether the magnitude of additive genetic, shared, and nonshared environmental influences on alcohol dependence symptoms varied as a function of decreasing age. Twin models were fitted in Mx. Results:, Risk for alcohol dependence symptoms increased with decreasing age at first drink. Heritable influences on alcohol dependence symptoms were considerably larger in those who reported an age at first drink prior to 13 years of age. In those with later onset of alcohol use, variance in alcohol dependence was largely attributable to nonshared environmental variance (and measurement error). This evidence for unmeasured gene × measured environment interaction persisted even when controlling for the genetic influences that overlapped between age at first drink and alcohol dependence symptoms. Conclusions:, Early age at first drink may facilitate the expression of genes associated with vulnerability to alcohol dependence symptoms. This is important to consider, not only from a public health standpoint, but also in future genomic studies of alcohol dependence. [source] Heritability of diurnal type: a nationwide study of 8753 adult twin pairsJOURNAL OF SLEEP RESEARCH, Issue 2 2007MARKKU KOSKENVUO Summary Twin studies suggest a genetic component in diurnal types. In 1981, a questionnaire sent to the Older Finnish Twin Cohort yielded responses from 2836 adult monozygotic (MZ) and 5917 like-sexed dizygotic (DZ) twin pairs with four category self-report on diurnal type. We used structural equation modelling to estimate genetic and environmental components of variance in morningness and eveningness. The model fitting was best when the morningness and the eveningness were analysed together. The ADE-model (including additive genetic, dominant genetic and non-shared environmental effects) fitted best to the data. ADE-models for men and women separately did not differ in a statistically significant manner from the combined model, and similarly ADE-models for young and old age groups separately did not differ either. The estimate for overall genetic effect (broad sense heritability) was 49.7% (95% confidence interval 46.4,52.8), with the remainder accounted for by environmental factors not shared by siblings. The variance component estimates for the underlying liability to diurnal type were 11.7% (95% CI 0,23.7) for additive genetic factors, 38.0% (24.7,51.3) for genetic factors due to dominance. Genetic effects thus account for about one-half of the interindividual variability in diurnal type in adults. [source] Stability of genetic influences on pulmonary function in a longitudinal study of octogenarian twinsAMERICAN JOURNAL OF HUMAN BIOLOGY, Issue 3 2010Terrie Vasilopoulos Using data from the first four waves of the OCTO-Twin study (twins 80 + years), the present study investigated the stability and change of genetic and environmental contributions to pulmonary function. Using a genetic simplex model, variance in peak expiratory flow (PEF) at each wave was decomposed into additive genetic and nonshared (specific) environmental factors. Additionally, this analysis distinguished the source of these influences, either from previous waves (transmissions) or from novel influences at each wave (innovations). At each time point (except wave 1), the genetic variance was due to genetic transmissions from prior time points. Conversely, the specific environmental variance in PEF at each time point was mainly due to environmental innovations. These results imply that genetic factors contribute to the stability of pulmonary function over time whereas environmental factors contribute to its change. Am. J. Hum. Biol., 2010. © 2009 Wiley-Liss, Inc. [source] Brief communication: Familial resemblance in digit ratio (2D:4D)AMERICAN JOURNAL OF PHYSICAL ANTHROPOLOGY, Issue 2 2009Martin Voracek Abstract Familial resemblance in the second-to-fourth digit ratio (2D:4D), a proxy for prenatal androgen action, was studied in 1,260 individuals from 235 Austrian families. In agreement with findings from twin studies of 2D:4D, heritability estimates based on parent,child and full-sib dyad similarity indicated substantial genetic contributions to trait expression (57% for right hand, 48% for left hand 2D:4D). Because twin studies have found nonadditive genetic as well as shared environmental effects on 2D:4D to be negligible or nil, these family-based estimates in all likelihood reflect the narrow-sense (additive genetic) heritability of the trait. Directional (right-minus-left) asymmetry in 2D:4D was only weakly heritable (6%). The pattern of same-sex and different-sex parent,child and full-sib correlations yielded no evidence for X-linked inheritance. This is surprising, considering evidence for associations of male 2D:4D with sensitivity to testosterone (functional variants of the X-linked androgen receptor gene). 2D:4D was particularly strongly heritable through male lines (father,son and brother,brother correlations), thus raising the possibility that Y-linked genes (such as the sex-determining region SRY) might influence 2D:4D expression. Am J Phys Anthropol, 2009. © 2009 Wiley-Liss, Inc. [source] Investigation of Gibbs sampling conditions to estimate variance components from Japanese Black carcass field dataANIMAL SCIENCE JOURNAL, Issue 5 2009Aisaku ARAKAWA ABSTRACT The genetic evaluation using the carcass field data in Japanese Black cattle has been carried out employing an animal model, implementing the restricted maximum likelihood (REML) estimation of additive genetic and residual variances. Because of rapidly increasing volumes of the official data sets and therefore larger memory spaces required, an alternative approach like the REML estimation could be useful. The purpose of this study was to investigate Gibbs sampling conditions for the single-trait variance component estimation using the carcass field data. As prior distributions, uniform and normal distributions and independent scaled inverted chi-square distributions were used for macro-environmental effects, breeding values, and the variance components, respectively. Using the data sets of different sizes, the influences of Gibbs chain length and thinning interval were investigated, after the burn-in period was determined using the coupling method. As would be expected, the chain lengths had obviously larger effects on the posterior means than those of thinning intervals. The posterior means calculated using every 10th sample from 90 000 of samples after 10 000 samples discarded as burn-in period were all considered to be reasonably comparable to the corresponding estimates by REML. [source] Heritability of Obesity-related Phenotypes and Association with Adiponectin Gene Polymorphisms in the Chinese National Twin RegistryANNALS OF HUMAN GENETICS, Issue 2 2010Juan Lee Summary The purpose of this study was to estimate the heritability of obesity-related phenotypes and investigate the association of adiponectin gene polymorphisms +45T>G and +276G>T with these measures in Chinese twins. 1260 twin pairs were recruited from two cities through the Chinese National Twin Registry System from 2001 to 2005. Two SNPs at the adiponectin locus (+45T>G and +276G>T) were genotyped. Structural equation modeling (SEM) was used to estimate heritability and the best-fitting variance component model. The regular association among all twins was analysed with generalised estimating equations (GEE). Sib-transmission/disequilibrium test (TDT) within dizygotic (DZ) twin pairs discordant for their genotype was performed using SEM. Additive genetic, common and unique environmental (ACE) model-based heritability of body mass index (BMI) was 61%, while additive genetic and unique environmental (AE)-model-based heritability of waist circumference (WC) and waist-hip ratio (WHR) were 75% and 61%, respectively. There was no association of adiponectin gene +45T>G and +276G>T genotypes with obesity-related phenotypes in all twins or discordant DZ twins. Our twins data did not support that there was an association between adiponectin gene polymorphisms +45T>G and +276G>T and the obesity-related phenotypes. Further studies are required to better understand the role of adiponectin gene polymorphisms in obesity. [source] Electrocardiographic Indices of Left Ventricular Hypertrophy and Repolarization Phase Share the Same Genetic Influences: A Twin StudyANNALS OF NONINVASIVE ELECTROCARDIOLOGY, Issue 4 2009Sara Mutikainen M.Sc. Background: Both left ventricular hypertrophy (LVH) and repolarization phase (RP) are known to be attributable to genetic influences, but less is known whether they share same genetic influences. The aim of this study was to investigate to what extent individual differences in electrocardiographic (ECG) LVH and RP are explained by genetic and environmental influences and whether these influences are shared between these two traits. Methods: Resting ECG recordings were obtained from 186 monozygotic and 203 dizygotic female twin individuals, aged 63 to 76 years. Latent factors, called LVH and RP, were formed to condense the information obtained from LVH indices (Cornell voltage and Cornell product) and T-wave amplitudes (leads V5 and II), respectively. Multivariate quantitative genetic modeling was used both to decompose the phenotypic variances into additive genetic, common environmental, and unique environmental influences, and for the calculation of genetic and environmental correlations between LVH and RP. Results: Additive genetic influences explained 16% of individual differences in LVH and 74% in RP. The remaining individual differences were explained by both common and unique environmental influences. The genetic correlation and unique environmental correlation between LVH and RP were ,0.93 and ,0.05, respectively. Conclusions: In older women without overt cardiac diseases, RP is under stronger genetic control than LVH. The majority of genetic influences are shared between LVH and RP whereas environmental influences are mainly specific to each. [source] Genetic Influences on Resting Electrocardiographic Variables in Older Women: A Twin StudyANNALS OF NONINVASIVE ELECTROCARDIOLOGY, Issue 1 2009Sara Mutikainen M.Sc. Background: 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. Methods: 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. Results: 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 TV5. The results also suggested dominance genetic influences on QRS duration, TV1, 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. Conclusion: 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. [source] Additive genetic and other sources of variation in growth traits of juvenile black bream Acanthopagrus butcheriAQUACULTURE RESEARCH, Issue 7 2005Robert G Doupé Abstract We used a factorial mating design to estimate the contribution of additive genetic, non-additive genetic and maternal effects to variation in growth traits of black bream Acanthopagrus butcheri (Munro) at 75, 130 and 180 days of age in the hatchery. Maternal genetic and environmental effects were greatest at 75 days of age, accounting for 9.1% of total phenotypic variance in wet weight, 11.4% of variance in standard length and 8.8% of variance in total length. At later ages maternal effects were much reduced, explaining 0.8,3.7% of phenotypic variance in growth traits. Additive genetic effects were greatest at 130 days of age, when they accounted for 17.4% of total phenotypic variance in wet weight, 21.4% of variance in standard length and 18.7% of variance in total length. Additive genetic effects were negligible (<1%) at 75 days of age and 4.8,5.5% of total phenotypic variance in growth traits at 180 days of age. Non-additive genetic effects (which also included common environmental effects because of families being raised in the same tank) explained 5.8,7.3% of total phenotypic variance in growth traits at 75 days of age, but were much smaller at later ages. Variable stocking densities among tanks up to 75 days significantly affected all growth trait measurements below 180 days of age. [source] Evolutionary genetics of genital size and lateral asymmetry in the earwig Euborellia plebeja (Dermaptera: Anisolabididae)BIOLOGICAL JOURNAL OF THE LINNEAN SOCIETY, Issue 1 2010YOSHITAKA KAMIMURA Male genitalia show several evolutionary characteristics, including rapid morphological divergence between closely related species and low within-species phenotypic variability. In addition, genital asymmetry is widespread despite the essentially bilaterally symmetric external morphology of insects. Several hypotheses, such as sexual selection and lock-and-key hypotheses, have been proposed to explain these characteristics of genital evolution. Although these hypotheses provide different predictions about the genetic basis of variation in genitalia, detailed quantitative genetic studies have been conducted in only three insect taxa: heteropterans, dung beetles (Scarabaeidae), and drosophilid flies. For an anisolabidid earwig, Euborellia plebeja, characterized by paired elongated intromittent organs, we estimated the heritabilities and genetic correlations of genital laterality, size of genitalia, and body size. No statistically significant additive genetic, dominance, maternal, or common environmental effects were detected for genital laterality (readiness to use either the left or the right intromittent organ). This result lends further support to the general rule that the direction of antisymmetric variations is randomly determined by non-genetic factors. Irrespective of the restricted phenotypic variation in genitalia compared with body size (allometric slope < 1), as observed in previous studies for other insects, these two traits showed a similar level of genetic variation, measured as the narrow sense heritability (h2) and the coefficient of additive genetic variation (CVA). Comparison suggests the causes of interspecific differences in genetic variability/correlation structures were developmental processes (holo- or hemimetabolous) and/or mode of sexual selection. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 101, 103,112. [source] | ||||||||||||||||