Selection,structure and the heritability of behaviour

Characters which are closely linked to fitness often have low heritabilities (V_A/V_P). Low heritabilities could be because of low additive genetic variation (V_A), that had been depleted by directional selection. Alternatively, low heritabilities may be caused by large residual variation (V_R=V_P – V_A) compounded at a disproportionately higher rate than V_A across integrated characters. Both hypotheses assume that each component of quantitative variation has an independent effect on heritability. However, V_A and V_R 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 26 ). 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 CV_A of domestic species. At the same time CV_R increased disproportionately faster than CV_A across integration levels, demonstrating variance compounding. However, neither CV_R nor CV_A had a predominant effect on heritability. The partial regression coefficients of CV_R and CV_A 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.     

X-CHROMOSOME INACTIVATION AND DEVELOPMENTAL PATTERNS IN MAMMALS

1. The review considers information from mammalian embryology relevant to X-chromosome inactivation, and from X-inactivation relevant to mammalian embryology. 2. Properties of the inactive-X, by which it may be recognized are: sex chromatin, heteropycnosis, late replication and the absence of gene product. Each of these has advantages and disadvantages in particular circumstances. In some species the X carries constitutive heterochromatin, which must be distinguished from the facultative region. 3. The time of X-chromosome inactivation can be estimated from the time of appearance of sex chromatin or late replication, or inferred from the appearance of heterozygotes for X-linked genes or of experimental chimaeras. The estimated time varies with species, and in the mouse and rabbit is near the time of increase in RNA synthesis. 4. Whereas in eutherian mammals either the maternally or the paternally derived X may be inactivated in different cell lines, in marsupials the paternal X is always the inactive one. 5. During development various factors act to distort the patterns produced by random X-inactivation. These factors include cell selection, transfer of gene product, and migration and mingling of cells. 6. There is no clear evidence that X-chromosome inactivation is not complete. 7. In female germ cells both X-chromosomes appear to be active. In male ones both X and Y appear inactive during most of spermatogenesis, although probably in early stages all X chromosomes present are active. 8. The active and inactive X-chromosomes may be differentiated by presence or absence of some non-histone protein or other polyanionic substance. 9. If the genes concerned in synthesis or attachment of this substance are on the X-chromosome then the differentiation will be self-maintaining. 10. The initiation of the differentiation requires either the attachment of different X-chromosomes to different sites, or some interaction of X-linked and autosomal genes, concerned in inducing or repressing activity. Some possible models are discussed.     

Mapping of the genes for dioecism and monoecism in Spinacia oleracea L.: evidence that both genes are closely linked

Spinach is basically a dioecious species, with occasional monoecious plants in some populations. Sexual dimorphism in dioecious spinach plants is controlled by an allelic pair termed X and Y located on the short arm of the longest chromosome (x = 6). Ten AFLP markers, closely linked to the X/Y locus, were identified using bulked segregant analysis, four of which were revealed to co-segregate with Y in the present mapping population. We mapped the AFLP markers and two known male-specific DNAs to a 13.4 cM region encompassing the locus. These markers will be the basis for positional cloning of the sex-determination gene. We also showed that a single, incompletely dominant gene is responsible for the highly staminate monoecious character. The gene was found to be located at a distance of 4.3 cM from microsatellite marker SO4, which mapped 1.6 cM from the X/Y locus. This indicates that the monoecious gene seems not to be allelic to but closely linked to the X/Y gene pair. SO4 will enable breeders to efficiently select highly male monoecious plants for preferential use as the pollen parent for hybrid seed production.     

Genetic and environmental variations in quantitative characters in fishes: a comparative analysis of monoclonal triploid and bisexual tetraploid spined loaches (<Emphasis Type="Italic">Cobitis</Emphasis>, Cobitidae)

The heritability estimates of 25 external morphometric characters and 23 craniometric indices are obtained by use of variances in monoclonal all-female triploids and bisexual tetraploids of spined loaches (genus Cobitis, Cobitidae) collected from the same breeding biotope. Most of studied traits demonstrate low heritability confirming previous conclusion on the similarity between external morphometric characters and craniological indices in relative effects of genetic and environmental components in their total phenotypic variation. Low heritability estimates in most of external morphological traits correspond to their low diagnostic value in Cobitis species. As a whole, in spite of certain deviations, studies on clonal forms do not refute the concept on higher heritability estimates in diagnostically significant traits in comparison with traits without diagnostic values in the same taxonomic group. Low heritability in most morphometric traits more probably is resulted from their low additive genetic variation caused by strong selection of evolutionary developed specific body shape in spined loaches, because strong selection should reduce the genetic variance in body proportions to minimal size. Sex differences observed in heritability estimates should be interpreted as a result of linkage of several additive genes controlling these traits to sex chromosomes. A few characters demonstrating high heritability estimates up to 0.492–0.580 are of great interest for taxonomic and phylogenetic studies in genus Cobitis and related taxa.     

Comparative Study of the Ribosomal RNA from Leishmania and Trypanosoma1

The ribosomal RNA from several stocks of the genera Leishmania and Trypanosoma were studied by gel electrophoresis, sedimentation on sucrose density gradients and RNA/DNA hybridization experiments. Three major components were observed after electrophoresis in polyacrylamide gels (PAGE-SDS), the relative molecular masses being respectively: X₁= 0.83 megadaltons, X₂= 0.63 megadaltons and X₃= 0.54 megadaltons for Leishmania RNA; and X₁= 0.86 megaldaltons, X₂= 0.78 megadaltons, and X₃= 0.58 megadaltons for Trypanosoma RNA. Depending upon the isolation procedure, a fourth component. X₀= 1.2 megadaltons (26S), became evident. The later component was purified from Leishmania brasiliensis (Y) by centrifugation on a linear 15-30% sucrose density gradient. This component, after heat denaturation and PAGE-SDS, gave rise to two bands coinciding in molecular mass with those of X₂ and X₃ indicating that these components are part of the large ribosomal subunit whereas X₁ belongs to the small one. The above mentioned differences in mobilities of components X₁ and X₂ between the two genera were no longer observed after electrophoresis in denaturing agarose-formaldehyde gels, suggesting secondary structural differences among these RNA species. Hybridization experiments with L. brasiliensis (Y) DNA showed that both RNA types compete equally well for the ribosomal sites in this DNA, and that L. brasiliensis (Y) rRNA recognizes the ribosomal sites in DNA of Trypanosoma cruzi (EP), thus indicating that no gross changes occurred in their nucleotide sequences during evolution.     

Sex determination and phenotype in wood lemmings with XXY and related karyotypic anomalies

Summary The wood lemming, Myopus schisticolor, possesses a unique sex determining system comprising both XX and XY females. Normal female development in the presence of XY is guaranteed by a mutation on the X, apparently associated with a structural rearrangement in Xp. This mutation inactivates the testis-inducing and male-determining factor on the Y and distinguishes X^* from X, and X^*Y females from XY males. Normal fertility of X^*Y females is ensured by a mitotic (double) nondisjunction mechanism which, at an early fetal stage, eliminates the Y from the germ line and replaces it by a copy of the X^*.Numerical sex chromosome aberrations are not infrequent and the trisomics XXY and X^*XY are relatively common. XXY individuals are sterile males with severe suppression of spermatogenesis. Among X^*XY animals, both males and females, as well as a true lateral hermaphrodite have been observed. Primary deficiency of germ cells, impairment of spermatogenesis and sterility are characteristic traits of the X^*XY males, whereas X^*XY females have normal oogenesis and are fertile. Both these extremes (except female fertility) coexist in the true hermaphrodite described in the present study. These apparently contradictory observations are explainable under the assumption that X^* and X in X^*XY individuals are inactivated non-randomly or that the cells are distributed unequally. Inactivation of the X or X^* determines whether or not the H-Y antigen will be expressed. When comparing conditions in Myopus and in man, an additional assumption has to be made in relation to the gene(s) involved in sex determination, located in Xp:In Myopus they do not escape inactivation, whereas in man they have been claimed to remain active.     

Characterizations of the Exponential Distribution by Local Maximum Random Variables

Let {X_n, n≧1} be a sequence of independent and identically distributed random variables. Suppose n(K)=min n for which X_n>X_k, with K fixed and n>K. We will characterize the exponential distribution by considering the distributional properties of X_k and X_k and X_n(k).     

On Encoding Values for Data Recorded as X ≥ C

In many medical and other contexts, an ordered variable is measured and the limits of measuring ability will be reached. In such cases, the data may be reported as X ≥ C. For example, in a cell count, a laboratory technician may report the data as “count ≥ 100.” If the data are a dependent variable, censored data methods are the standard techniques. However, if X is an independent variable, some methods need to be devised to handle the problem. A simple method is to encode the values X ≥ C to some value X₀. A candidate for this is the conditional mean (or median) of X given X ≥ C. This paper gives some results, and examines some robustness questions for the proposed method.     

Twin‐based heritability of actimetry traits

There is a critical need for phenotypes with substantial heritability that can be used as endophenotypes in behavioral genetic studies. Activity monitoring, called actimetry, has potential as a means of assessing sleep and circadian rhythm traits that could serve as endophenotypes relevant to a range of psychopathologies. This study examined a range of actimetry traits for heritability using a classic twin design. The sample consisted of 195 subjects from 45 monozygotic (MZ) and 50 dizygotic (DZ) twin pairs aged 16‐40 years. Subjects wore both a research‐grade actimeter (GENEActiv) and a consumer‐oriented device (FitBit) for 2 weeks. Sleep and circadian traits were extracted from GENEActiv data using PennZzz and ChronoSapiens software programs. Sleep statistics for a limited number of FitBit‐collected traits were generated by its accompanying mobile app. Broad sense heritability was computed on a set of 33 MZ and 38 DZ twin pairs with complete data using both OpenMX and SOLAR software. These analyses yielded a large number of actimetry‐derived traits, 20 of which showed high heritability (h² > 0.6), seven of which remain significant after Bonferroni correction. These results indicate that actimetry enables assessing a range of phenotypes with substantial heritability that may be useful as endophenotypes for genetic studies.     

Cytological identification of two X-chromosome types in the wood lemming (Myopus schisticolor)

In the wood lemming (Myopus schisticolor) three genetic types of sex chromosome constitution in females are postulated: XX, X^*X and X^*Y (X^*=X with a mutation inactivating the male determining effect of the Y chromosome). Males are all XY. It is shown in the present paper that the two types of X chromosomes, X and X^*, exhibit differences in the G-band patterns of their short arms. In addition, it was demonstrated in unbanded chromosomes that the short arm in X^* is shorter than in X. The origin of these differences is still obscure; but they allow to identify and to distinguish the individual types of sex chromosome constitution, as of XX versus X^*X females and of X^*Y females versus XY males, on the basis of G-banded chromosome preparations from somatic cells.     

Repeatability: Its role in evolutionary studies of mating behavior

Summary Repeatability, a concept derived from quantitative genetics theory, is a statistic that describes the degree to which variation within individuals contributes to total variation in a population. Its usual application has been to set an upper limit on heritability but it may also be useful for studies of stereotypy of behavior. The repeatability of the production of male mating signals gives information both about whether males differ sufficiently for selection to act and whether the differences could be appreciably heritable. Measures of the repeatability of female mating preferences will provide data that can describe the preference functions used in mathematical models of the evolution of sexually selected traits, as well as putting an upper bound on the heritability of preferences. A survey of the few measures in the literature shows that the repeatability of male signal production varies substantially (range 0.21–0.85) and does not necessarily reflect heritability. The repeatabilities of female preferences have not been published previously: for the response to conspecific pheromones by female flour beetles (Tribolium castaneum), my best estimate is zero. Measuring the repeatability of other traits such as parental care and foraging behavior may also lead to insights about selection on and the evolution of these traits.     

Cytotaxonomy in distinct populations of Hoplias aff. malabaricus (Characiformes,Erythrinidae) from lower Paranapanema River basin

Cytogenetic and morphometric analyses were carried out in Hoplias aff. malabaricus specimens from six distinct populations from the lower Paranapanema River basin, located between the states of Paraná and São Paulo, Brazil. Measurements were also taken from a specimen collected in Surinam. In a population from a fish farm station at Universidade do Norte do Parana (EPUNOPAR), two sympatric cytotypes (2n = 40 and 2n = 42 chromosomes) are found. A population from a fish farm station at Universidade Estadual de Londrina (EPUEL) shows 2n = 42 meta–submetacentric chromosomes for males and females with a simple sex chromosome system of XX/XY type. Populations from the Vermelho and Rancho Alegre Rivers, Três Bocas Stream and Paranapanema River have 2n = 39 chromosomes in males and 2n = 40 chromosomes in females, showing a multiple sex chromosome system of X₁X₁X₂X₂/X₁X₂Y type. Twenty morphological variables were studied. These measurements were used for an analysis of the canonical variables and standard analysis of proportional measurements. The most variable measurements among the specimens are the maxilla length (MXL) and the pre‐dorsal distance (PDD). Analysis of canonical variables indicates three distinct groups in the first canonical axis formed by: (1) Três Bocas Stream, (2) Rancho Alegre + Vermelho River + EPUNOPAR and (3) EPUEL + Paranapanema River. This axis retained 79·4% of information from the original matrix. Analysis of morphometrics reveals differences among populations from the Paranapanema River basin and between these and the specimen from Surinam. The morphometric and cytogenetic differences among the studied populations suggest a species complex.     

Effects on ecosystem resilience of biodiversity, extinctions, and the structure of regional species pools

Resilience is a general concept that aims to help understand how ecosystems respond to disturbances such as extinctions and invasions. Here, we propose a measure of one aspect of resilience, R _X , which is one minus the expected change in functional diversity (X) caused by a species extinction or addition. We show how two components of biodiversity, species richness and functional diversity, and the structure of regional species pools affect this measure. Variation in species richness and in functional diversity have opposite effects on R _X . Speciose assemblages generally have higher R _X than depauperate ones, whereas functionally diverse assemblages have low R _X relative to functionally depauperate ones. The effect of an extinction on R _X reflects this tradeoff. In our analyses, extinctions usually cause only a small decrease in both functional diversity and R _X . However, extinctions sometimes cause a large reduction in functional diversity and then tend to increase R _X . Regional assemblages containing all rather unique species tend to result in speciose assemblages with relatively low R _X and in low richness assemblages with relatively high R _X . The opposite is true of regional assemblages containing functionally similar species. Information about the processes that structure regional assemblages will therefore increase understanding of ecosystem resilience. Generally, these results suggest that management for biodiversity may not always result in management for resilience.     

Evaluation of the Fermi equation as a model of dose-response curves

 When plotted in linear coordinates, the dose-response curves of microorganisms exposed to a lethal agent, such as radiation or a toxic substance, often have a characteristic sigmoid shape. Irrespective of whether they are very narrow or broad they can be described by the Fermi function, which is a mirror image of the logistic function, i.e. S(X)=1/{1+ exp [(X−X _c)/a]} where S(X) is the fraction of the surviving organisms, X the dose of the lethal agent, X _c a characteristic dose marking the inflection point of S(X), which corresponds to 50% mortality, and a a measure of the steepness of the survival curve around X _c. It is demonstrated that, if the susceptibilities of the individual organisms, expressed in terms of a characteristic lethal dose, have a symmetric unimodal distribution, the dose-response curve of the population has a Fermian sigmoid shape. It is also shown that the mode and variance of the distribution can be estimated from the shape parameters of the Fermian survival curve, X _c and a. Received: 7 November 1995 / Received last revision: 11 April 1996 / Accepted: 29 April 1996     

Evaluation of circulant partial diallel crosses in maize

Summary The present study was conducted in maize (Zea mays L.) on crosses among 20 diverse parents. The materials were evaluated in four different environments for eight characters. Combining ability analysis was carried out following diallel and partial diallel crosses. The number of crosses per parent (s) varied from 3 to 19 and the results were studied to identify the critical value of s that would provide an adequate information with minimum resources. The S₅ partial diallel was as good as the S₁₉ for the detection of differences among general combining ability (GCA) effects. Even the S₃ gave adequate information in the case of characters with high heritability. However, partial diallel analysis was less efficient in detecting the differences due to specific combining ability (SCA) effects. These results varied with environments, and characters with low heritability were more prone to misinterpretation. GCA effects showed fluctuations in partial diallel analysis which were more pronounced in S₅ and S₃, particularly for characters with low heritability. The average standard error of difference between GCA effects increased with a decrease in s, with a steep slope for s < 7. The partial diallel analysis was more efficient for the estimation of the variance component of GCA than for SCA, as the estimate of SCA was biased upwards. Estimates of broad sense heritability obtained from the partial diallels agreed with the full diallel analysis better than the narrow sense estimates. Smaller partial diallels gave erratic estimates of heritability, particularly for the characters with low heritability.     

SYNTHETIC AGROPYRON-ELYMUS HYBRIDS: III. ELYMUS CANADENSIS X AGROPYRON CANINUM,A. TRACHYCAULUM,AND A. STRIATUM

Hybrids were produced with relative ease from controlled crosses of Elymus canadensis L. with European Agropyron caninum (L.) Beauv., North American A. trachycaulum (Link) Malte ex H. F. Lewis, and Asian A. striatum Nees ex Steud. All hybrids appeared to be completely sterile and were, for the most part, morphologically intermediate between their parents. The E. canadensis × A. caninum hybrids were exceptionally vigorous and leafy and may have some potential as forage grasses if fertility can be achieved. All parent plants were tetraploid, 2n = 28, and they behaved cytologically as alloploids. Chromosome pairing in the hybrids indicated that both E. canadensis genomes were closely homologous with those of A. trachycaulum and somewhat less homologous with those of A. caninum. Interchanged and inverted chromosome segments apparently constitute the major differences between E. canadensis, A. trachycaulum, and A. caninum genomes; however, cryptic structural differences must also exist. Partial homologies were detected between one A. striatum and E. canadensis genome, but their other genomes were distinctly different. The genome relations between the parent species were expressed in terms of the following genome formulas: E. canadensis, S₁S₁X₁X₁; A. trachycaulum, S₂S₂X₂X₂; A. caninum, S₃S₃X₃X₃ : and A. striatum S₄S₄YY or X₄X₄YY, where “S” refers to a genome derived from A. spicatum and “X” and “Y” are genomes of unknown origin.     

Resemblance for Body Mass Index in Families of Obese African American and European American Women

Objective: We determined the levels of resemblance in body mass index (BMI) in large samples of families selected through obese African American and European American women. Research Methods and Procedures: We examined correlations among relatives in 1185 European American and African American families ascertained through age-matched obese women (BMI ≥ 30 kg/m²). A subset of 801 families were ascertained through extremely obese women (BMI ≥ 40 kg/m²). Results: Parent-offspring and sibling correlations ranged from 0.19 to 0.15, suggesting a moderate level of heritability in both groups. Mean BMI values for female relatives were lower for European Americans than for African Americans even though probands were matched, perhaps because the European American relatives regress to a lower population mean. We found significantly higher family correlations for height in European Americans, suggesting greater environmental variability among African Americans for factors affecting growth and physical development. Discussion: Our results suggest a similar level of heritability of BMI in families of obese African American and European American women. Other genetic studies will be needed to determine the extent to which the same or different genes and environmental conditions contribute to an overall similar heritability in the two racial groups.     

Estimating overdispersion in sparse multinomial data

Multinomial data arise in many areas of the life sciences, such as mark-recapture studies and phylogenetics, and will often by overdispersed, with the variance being higher than predicted by a multinomial model. The quasi-likelihood approach to modeling this overdispersion involves the assumption that the variance is proportional to that specified by the multinomial model. As this approach does not require specification of the full distribution of the response variable, it can be more robust than fitting a Dirichlet-multinomial model or adding a random effect to the linear predictor. Estimation of the amount of overdispersion is often based on Pearson's statistic X² or the deviance D. For many types of study, such as mark-recapture, the data will be sparse. The estimator based on X² can then be highly variable, and that based on D can have a large negative bias. We derive a new estimator, which has a smaller asymptotic variance than that based on X², the difference being most marked for sparse data. We illustrate the numerical difference between the three estimators using a mark-recapture study of swifts and compare their performance via a simulation study. The new estimator has the lowest root mean squared error across a range of scenarios, especially when the data are very sparse.