RESISTANCE OF GENETIC CORRELATION STRUCTURE TO DIRECTIONAL SELECTION IN DROSOPHILA MELANOGASTER

The genetic covariance and correlation matrices for five morphological traits were estimated from four populations of fruit flies, Drosophila melanogaster, to measure the extent of change in genetic covariances as a result of directional selection. Two of the populations were derived from lines that had undergone selection for large or small thorax length over the preceding 23 generations. A third population was constituted using flies from control lines that were maintained with equivalent population sizes as the selected lines. The fourth population contained flies from the original cage population from which the selected and control lines had been started. Tests of the homogeneity of covariance matrices using maximum likelihood techniques revealed significant changes in covariance structure among the selected lines. Prediction of base population trait means from selected line means under the assumption of constant genetic covariances indicated that genetic covariances for the small population differed more from the base population than did the covariances for the large population. The predicted small population means diverged farther from the expected means because the additive genetic variance associated with several traits increased in value and most of the genetic covariances associated with one trait changed in sign. These results illustrate that genetic covariances may remain nearly constant in some situations while changing markedly in others. Possible developmental reasons for the genetic changes are discussed.     

Genetic covariances within and between species: indirect selection for hybrid inviability

Genotype-environment interactions and natural selection can result in local specialization when different genotypes are favored in different environments. Restricted gene flow or genetic subdivision enhances local genetic diversification across a species when natural selection acts on such variation. The indirect evolution of reproductive isolation and the restriction of gene flow between species in statu nascendi may provide a central role for genotype-environment interactions in speciation genetics. We derive the expected genetic covariance between heterospecific and conspecific viability fitness under several different models of selection, dominance, and breeding structure. Standard quantitative genetic methods can be used to estimate these covariances in experimental studies. These genetic covariances permit us to evaluate in a formal way the indirect effects of selection within a species on the evolution of hybrid inviability between species. We find that, for autosomal loci and random mating, the genetic covariance across species is equal to the product of three quantities: (1) the viability of the best hybrid genotype; (2) the viability effect of an allele in hybrids; and, (3) the change in allele frequency due to selection in the conspecific population. Inbreeding within the conspecific population, expressed as Wright's coefficient, F, increases the genetic covariance by a factor (1 + F). In all cases, a negative genetic covariance across species is evidence for hybrid inviability evolving as an indirect effect of selection within species for adaptive (as opposed to neutral) genetic change. “It is an irony of evolutionary genetics, that although it is a fusion of Mendelism and Darwinism, it has made no direct contribution to what Darwin obviously saw as the fundamental problem: the origin of species…. While it is a question of elementary population genetics to state how many generations will be required for the frequency of an allele to change from q₁ to q₂, we do not know how to incorporate such a statement into speciation theory, in large part because we know virtually nothing about the genetic changes that occur in species formation.” (Lewontin 1974, p. 159)     

Doubled Haploids for Studying the Inheritance of Quantitative Characters

By using a doubled-haploid population derived from F₂ plants, additive and additive x additive genetic variances, as well as the number of segregating genes, can be estimated. An F₂-derived doubled-haploid population may contain almost 50% more of the best recombinant than an F₁-derived population. However, the best recombinant occurs in the same frequency in the two populations when there is no linkage between genes. The difference in the frequency of the best recombinant between F₂- and F₃-derived populations is small. This implies that the doubled-haploid method using F₂ plants provides only slightly less opportunity for recombination than the conventional breeding methods of self-pollinating crops. In the absence of additive epistasis, a weighted mean of recombination values can be estimated using an F₂-derived population and its parental lines. When additive epistasis is present, it can be estimated from doubled-haploid populations derived from two backcrosses. Studies on the linkage of quantitative characters are needed for determining whether doubled haploids should be produced from F₂ or from F₁ plants in a breeding program.     

Comparison of bread wheat lines selected by doubled haploid, single-seed descent and pedigree selection methods

 Yield performance of each group of ten spring bread wheat lines selected by doubled haploid (DH), single-seed descent (SSD) and pedigree selection (PS) methods from three F₁ crosses was compared with the aim of evaluating the DH method in breeding programs. Populations of 65–97 DH lines and 110 SSD lines per cross were used for selection. PS lines were developed by repeated selections from 1500 F₂ plants. Yield evaluation was performed at the F₆ generation of SSD and PS lines along with DH lines in a 2-year field experiment. It took only 2 years from the planting of wheat materials for DH production to the planting of selected DH lines for yield evaluation. There was no significant difference in grain yield between DH lines and PS lines selected from an F₁ cross whose parental varieties were closely related in their pedigrees. In two crosses with low coefficients of parentage and a large variation in their progenies, grain yield of selected DH lines was significantly lower than those of selected SSD and PS lines. These results confirm that the DH method can save time in obtaining recombinant inbred lines ready for yield evaluation. However, a larger DH population is required to achieve the same level of genetic advance with the PS method in crosses containing greater genetic variation. Received: 23 December 1997 / Accepted: 12 March 1998     

Effect of selection on the heterozygosity of inbred lines of maize

Although pedigree selection is the most commonly used method for developing inbred lines of maize, there are no studies on its effect on the heterozygosity of the lines. The objective of this work was to study the effect of pedigree selection on their heterozygosity. Thirteen F₅ or F₆ maize inbred lines developed by the pedigree selection method in four breeding programs and their F₁ and F₂ − F₄ ancestors were genotyped with simple sequence repeat markers distributed along the genome. Simulation was also conducted assuming different models of selection to investigate the selective forces needed to explain the data. In the F₂, F₃ and F₄ 40%, 66% and 86% of the markers segregating in the F₁ were fixed; that is, in the F₂ and F₃ fixation was lower than neutral expectation, but higher in the F₄. Due to such opposite apparent directions of selection, the heterozygosity of the lines in the F₅ or F₆ generations did not differ significantly from neutral expectations. The time to fixation differed from that expected with neutrality for most of the chromosomes, indicating that selection is distributed across the genome; but apparent overdominant effects in chromosome 7 were higher than in other chromosomes. In conclusion, the relationship between heterozygosity and vigour may reduce the effectiveness of pedigree selection in its goal of selecting the more vigorous, homozygous individuals. A more effective procedure is proposed using molecular markers for the identification of the more homozygous individuals, the most vigorous of those individuals being selected.     

DNA divergence as a criterion of a sweet pepper (Capsicum annuum L.) selection for heterosis

The identification of perspective parental lines for the creation of high-yield hybrids is the most labor-consuming stage of selection, because it needs extensive trials of combining ability. Based on evaluation of the genetic divergence of the parental lines, the prediction accuracy of F₁ hybrids performance was investigated. The value of the divergence was calculated using biometric and molecular methods, such as inter simple sequence repeats (ISSR) and random amplified polymorphic DNA (RAPD). Based on estimates of divergence, 10 lines were chosen for cyclic cross (scheme I) and testcross (scheme II). In most crosses, the F₁ hybrids were significantly superior to the parents in the main economically valuable traits. The level of heterosis was higher among hybrids of scheme I. Analysis of the relationship between parental divergence and F₁ performance showed that the hybrid productivity of scheme I was predetermined by ISSR divergence in 86%, and productivity was caused by RAPD divergence in 69%, whereas the F₁ yield of scheme II was not related to the value of genetic distances. Since the values of DNA divergence were closely associated both with midparent level and F₁ performance, we assumed that part of the polymorphic DNA fragments of the parental lines of scheme I is related to heterotic loci (HTL), which may be considered potential key markers for the heterotic selection of the sweet pepper.     

North American study on essential derivation in maize: inbreds developed without and with selection from F2 populations

An essentially derived variety largely retains the characteristics of a parental or ancestral variety. A consensus has not been reached regarding the threshold for declaring essential derivation in maize (Zea mays L.), partly because benchmark data are lacking. Our objective in this study, commissioned by the American Seed Trade Association, was to determine the range of parental contribution among maize inbreds developed without and with selection. Seed companies in North America contributed existing proprietary data on the molecular marker similarity of 100 or more families, developed without selection from F₂ populations, with each of their parents. The companies also sent us seed samples of elite inbreds, developed with selection from F₂ populations, for analysis using 60 RFLP marker loci and 20 SSR marker loci. Among the families developed without selection, the average parental contribution was close to the expected value of 0.50 for F₂ populations. Specific families received up to 79% of their alleles from one parent. Although selection tended to increase the frequency of such transgressive segregants, it did not necessarily increase the maximum parental contribution in an F₂ population. Parental contributions were consistent between the elite inbreds and their early-generation families. We conclude that inbreds with 70% to nearly 80% of their genome derived from one parent can be obtained from an F₂ population. Further empirical data would be valuable particularly for backcross populations, which were not available in this study. Received: 21 June 2000 / Accepted: 28 July 2000     

Prediction of hybrid grain yield performances in Indica Rice (<Emphasis Type="Italic">Oryza sativa</Emphasis> L.) with effect-increasing loci

One hundred and fifty-one rice hybrids produced in two sets of half-dialell crosses and their parents (13 cytoplasmic male sterile lines and 19 restorers) were used to predict the F₁ performances of seven yield traits through the parental genetic distances (GD) based on SSR markers. The positive loci (PL) and effect-increasing loci (IL), which were screened from SSR polymorphic loci by the F₁ traits of 32 parents, together with total loci (TL), were utilized to estimate parental GD and the models were found to predict the traits of hybrids derived from different parents, fixed parents, and different environments, respectively. The results were as follows: (1) 550 polymorphic loci were detected from 174 SSR markers: a dendrogram based on these loci could separate all the sterile and restorer lines used in the present study, which indicated that parental genetic diversity of F₁ was large; (2) the correlations between F₁ traits and parental GDs based on IL ranged from 0.61 to 0.87 with a mean of 0.76, and they were higher than those on TL or on PL; (3) predictions based on IL for F₁ traits (except grain weight per plant) derived from different environments were ideal, but worse for F₁ traits derived from different parents; and (4) IL was more effective than TL and PL in predicting traits of F₁ with fixed parents, and predictions for fixed restorer combinations were more effective than those for fixed sterile line combinations. These results should facilitate molecular prediction for hybrid yield and other traits by means of both elite sterile and restorer lines.     

Genotype-environment interaction effects on reproductive performance in Tribolium castaneum

Summary The genetical expectations of the means, variances and covariances of populations of doubled haploid lines derived from f₁, F₂, F₃ and intermated F₂ (S₃) generations are presented. These expectations are identical, regardless of genetical architecture, providing there is no linkage disequilibrium. In the presence of linkage disequilibrium differences will occur whose magnitude and direction will depend on the degree of disequilibrium, recombination frequency and the presence or absence of epistasis.Data from an experiment to detect linkage disequilibrium in a cross between two spring barley varieties are presented. This involved a comparison of means, variances and covariances of doubled haploid populations derived from the F₁ and F₂ generations using the H. bulbosum system. Linkage disequilibrium was detected for important agronomic characters and the effect of this disequilibrium on the choice of generation for doubled haploid production is discussed.     

Rapid inbreeding in maize

The number of inbreeding generations required to produce homozygotes may be reduced if the more homozygous individuals in each breeding generation are selected phenotypically in the segregating progenies for further inbreeding. To demonstrate this, inbred lines and their F₁, S₁, and S₂ progenies were studied for number of days from planting to anthesis, plant height and total leaf number. The data indicated that a high degree of control of rate of inbreeding may be obtained by proper use of heterotic characteristics in selection. The technique is a simple one: the selection of those individuals in a segregating array that most closely approach the characteristics of the ultimate homozygotes obtainable from the population. If these traits are undefined, then simple negative selection for heterotic attributes should suffice. In an S₁ progeny of maize, an individual with the same number of leaves and (1) as short as and (2) as slow to flower as the parental inbreds will be likely to be more homozygous than its taller and earlier flowering sibs of like leaf number. Since the inbreeder’s aim is not homozygosis per se but the development of agronomically useful homozygotes, the population dealt with should be sufficiently large to permit intense selection for outstanding individuals among the more homozygous members of each generation of selection.     

Genetic variability and advance under four selection procedures in wheat pedigree breeding programme

Summary Four methods of generation advance (SPS, SSD, BP and MMS) were compared in F₃ and F₄ generations. In the F₃ generation, the SPS and SSD methods of generation advance proved superior to the BP and MMS methods for grain yield per plant and for at least one of the yield component traits. The F₃ SSD population did not differ significantly from the F₃ SPS for any of the traits. However, the F₃ SSD population retained more range and cv for different traits than with other methods of generation advance. F₄ progenies derived from F₃ SSD population were significantly superior for grain yield than lines derived from the other three F₃ populations. The MMS method of generation advance proved useful for increasing the 1,000-grain weight for which initial selection was made.Part of Ph.D. Thesis submitted by senior author to Haryana Agricultural University, Hisar     

Should maize doubled haploids be induced among F<Subscript>1</Subscript> or F<Subscript>2</Subscript> plants?

Maize (Zea mays L.) doubled haploid lines are typically produced from F₁ plants. Studies have suggested that the low frequency of recombinants in doubled haploids may reduce the response to selection. My objective was to determine if, for sustaining long-term response, doubled haploids should be induced in F₁ or F₂ plants during maize inbred development. In simulation experiments, I examined the response to multiple cycles of testcross selection among doubled haploid lines derived from F₁ plants (denoted by DH), doubled haploid lines derived from F₂ plants (DH_F2), and recombinant inbred (RI) lines derived by single-seed descent. For a trait controlled by 100 or more quantitative trait loci (QTL), the cumulative responses to selection were up to 4–6% larger among DH_F2 lines than among DH lines. The cumulative responses were up to 5–8% larger among RI lines than among DH lines. The QTL become unlinked as the number of QTL in a finite genome decreases, and the responses among RI, DH, and DH_F2 lines were equal or nearly equal when only 20 QTL controlled the trait. Metabolic-flux epistasis reduced the differences in the response among RI, DH, and DH_F2 lines. Overall, the results indicated that doubled haploids should be induced from F₂ plants rather than from F₁ plants. If year-round nurseries are used and new F₁ crosses for inbred development are initially created on a speculative basis, the development of doubled haploids from F₂ rather than F₁ plants should not cause a delay in inbred development.     

Similarity of maize and sorghum genomes as revealed by maize RFLP probes

Summary Densely saturated genetic maps of neutral genetic markers are a prerequisite either for plant breeding programs to improve quantitative traits in crops or for evolutionary studies. cDNA and genomic clones from maize were utilized to initiate the construction of a RFLP linkage map in Sorghum bicolor. To this purpose, an F₂ population was produced from starting parental lines IS 18729 (USA) and IS 24756 (Nigeria) that were differentiated with regard to many morphological and agronomical traits. A total of 159 maize clones were hybridized to the genomic DNA of the two parents in order to detect polymorphism: 154 probes hybridized to sorghum and 58 out of these were polymorphic. In almost all of the cases hybridization patterns were similar between maize and sorghum. The analysis of the segregation of 35 polymorphic clones in an F₂ population of 149 individuals yielded five linkage groups. The three principal ones recall regions of maize chromosomes 1, 3 and 5: in general, colinearity was maintained. A possible inversion, involving a long region of maize chromosome 3, was detected. Simulations were also performed to empirically obtain a value for the lowest number of individuals of the F₂ population needed to obtain the same linkage data.Prof. E. Ottaviano, to whom this paper is dedicated, suddenly died on June 7^th, 1991     

HYBRIDIZATION IN LEOPARD FROGS (RANA PIPIENS COMPLEX): LARVAL FITNESS COMPONENTS IN SINGLE-GENOTYPE POPULATIONS AND MIXTURES

Recognizing the predominant mode of selection in hybrid systems is important in predicting the evolutionary fate of recombinant genotypes. Natural selection is endogenous if hybrid genotypes are at a disadvantage relative to parental species independent of environment. Alternatively, relative fitness can vary in response to environmental variation (exogenous selection), and hybrid genotypes can possess fitness values equal to or greater than that of parental species. I investigated the nature of natural selection in a leopard frog hybrid system by rearing larvae of hybrid and parental genotypes between Rana blairi and R. sphenocephala in 1000-L outdoor experimental ponds. Three hybrid (F₁, backcrossj [B₁], backcross₂ [B₂]) and two parental (R. blairi [BB] and R. sphenocephala [SS]) larval genotypes were produced by artificial fertilzations using adult frogs from a natural population in central Missouri. Resultant larvae were reared in single-genotype populations and two-way mixtures at equal total numbers from hatching to metamorphosis. In single-genotype ponds, F₁ hybrid larvae had highest survival and BB were largest at metamorphosis. When F₁ and SS larvae were mixed together, F₁ hybrids had reduced survival and both F₁ and SS larvae metamorphosed at larger body masses than when reared separately. When mixed, both B₁ and SS larvae had shorter larval period lengths than when reared alone. Higher proportion of B₁ metamorphs were produced when larvae were mixed with either parental species than when reared alone. Larval fitness components as measured by survival, body mass at metamorphosis, proportion of survivors metamorphosing, and larval period length for B₂ hybrid and BB larvae were similar in single-genotype populations and mixtures. Comparison of composite fitness component estimates indicated hybrid genotypes possess equivalent or higher larval fitness relative to both parental species for the life-history fitness components measured. Despite reduced survival of F₁ hybrids in mixtures, backcross-generation hybrid genotypes demonstrated high levels of larval growth, survival, and metamorphosis in mixtures with parental species. Consequently, this study suggests natural hybridization and subsequent backcrossing between R. blairi and R. sphenocephala can produce novel and relatively fit hybrid genotypes capable of successful existence with parental species larvae. Thus, the evolutionary fate of hybrid and parental genotypes in this system may be influenced by exogenous selection mediated by genotypic composition of larval assemblages.     

Cryptic species in Diacyclops bicuspidatus (Copepoda:Cyclopoida): evidence from crossbreeding studies

The problem of cryptic species in Diacyclops bicuspidatus was examined using interpopulation crosses of four populations collected from a: (1) permanent flood lake in Kiev, Ukraine, (2) temporary pool in Kiev, (3) permanent pond in St. Petersburg, Russia (1200 km to north from Kiev) and (4) lake in Crimea (1100 km south of Kiev). The only interpopulation crosses to exhibit fertility were those between the St. Petersburg population and each of the two Kiev populations. The crosses between the Kiev and Crimea populations, between the St. Petersburg and Crimea populations, and between the two Kiev populations were sterile, as evidenced by either nonviable eggs, empty egg membranes or incomplete copulations. The F₁ hybrids resulting from the St. Petersburg permanent pond X Kiev flood lake cross were fertile and produced mature F₂ offspring. Some data on development times of parental and hybrid lines are presented. The St. Petersburg parental line showed development times almost twice as long as those of the Kiev flood lake population when reared at 10 °C and 20 °C in the laboratory. The F₁ offspring of the cross between St. Petersburg females and Kiev floodlake males showed similar development times to females of the St. Petersburg parental lines at both temperatures. The F₂ hybrids also showed development times that approximated those of the St. Petersburg parental line. These crossbreeding studies suggest the presence of cryptic species in the D. bicuspidatus inhabiting ecologically different populations in many parts of its large holarctic range.     

Effects of different base populations on selection response in Drosophila

Of two laboratory strains of Drosophila melanogaster used in this study, the +3 strain had slightly higher mean abdominal bristle number and estimated heritability of this character than the Oregon‐R. Their F₁ hybrid exhibited 5 % heterosis. Fourteen generations of the two original strains and the F₃ of the hybrid were selected for high and low numbers of abdominal bristles on the 4th and 5th sternites, at a selection intensity of 20%. A mass‐mated unselected control was maintained for each population. The +3 population responded considerably more to selection for low numbers of bristles than high, and the Oregon‐R population showed a similar, though less marked, tendency; the Crossbred population responded more strongly to selection for high numbers. Except for the Crossbred high selection line, all lines declined in response rate, phenotypic variance, and realised heritability. The average realised heritability of the Oregon‐R and +3 high and low selection lines over 14 selection generations fell short of their predicted base population heritabilities. The deviation from the predicted was particularly pronounced with selection for high bristle number in the +3 line.     

Utilization of leaf temperature for the selection of leaf gas-exchange traits to induce heat resistance in sunflower (Helianthus annuus L.)

Heat stress is a major production constraint of sunflower worldwide. Therefore, various populations (parental, F₁, F₂, F₃, and plant progenies) of sunflower were screened for leaf gas-exchange traits with the objectives to formulate selection criteria of heat resistance and development of heat-resistant lines. Initial screening and F₂ seeds exposed to heat stress (45°C) resulted in the development of an adapted F₂ population that showed leaf gas-exchange and morphological traits better than the unadapted population. Correlation coefficients of traits were partitioned into direct and indirect effects via a path analysis technique to determine the cause of their relationship with a basic parameter such as a reproductive head mass (HM). Path analysis showed a positive direct effect of leaf temperature (T_leaf) (0.32) on HM and also an indirect effect (0.77) of the transpiration rate (E) on HM. Moreover, T_leaf showed high heritability estimates. T_leaf was used to select superior plants within the F₂ population. This selection brought about an improvement in the net photosynthetic rate (P _N) and E as it was indicated from progeny performance and realized heritability. Progenies selected on the basis of T_leaf also showed an increase in achene yield and heat resistance over unselected F₃ progenies and a commercial hybrid.     

Genetically idiosyncratic responses of Drosophila melanogaster populations to selection for improved learning ability

To what extent is adaptive evolution over short timescales repeatable? To address this question, we studied the performance of crosses between replicate Drosophila melanogaster lines previously subject to selection for improved learning response in the context of oviposition substrate choice. Of the 10 pairwise F₁ crosses among the five selection lines, four performed in the original learning assay similarly to the parental lines, whereas the remaining six showed learning scores significantly below the average of the parental lines. In particular, four F₁ crosses (three involving the same line) showed no detectable learning, on a par with unselected control lines. This indicates that the response to selection in some lines involved allelic substitutions at different loci. Additional assays of crosses between two selection lines indicated that the loss of performance in hybrids generalized to another type of learning assay, and held for both short‐ and long‐term memory. Joint analysis of first‐ and second‐generation crosses between these two lines supported the hypothesis that the response to selection in these different lines was based on the spread of recessive alleles at different loci. These results show that the evolutionary trajectories of populations of the same origin subject to uniform selection may sometimes diverge over very short evolutionary timescales.     

Morphological differentiation following experimental bottlenecks in the butterfly Bicyclus anynana (Nymphalidae)

The consequences of population bottlenecks for morphological differentiation were investigated experimentally in the Afrotropical butterfly Bicyclus anynana (Satyrinae). A genetically variable laboratory population was used to establish daughter populations differing in the severity of the founding bottleneck (two, six, 20 and c. 300 individuals), with four to six replicate lines per treatment. Nine quantitative traits of the hind wing were measured (wing area and eight wing pattern characters) in the founders, F₁, F₂ and F₃ generations. A tenth character, egg weight, was measured in all lines in F₃. The validity of the neutral additive model was tested by regressing the observed phenotypic variance among replicate lines (V_d) at F₃ against the expected additive genetic variance among lines (2F_tV_A0, where F_t is the coefficient of inbreeding at generation t and V_A0 is the additive genetic variance in the base population). This analysis was performed for the first six principal components of the wing character set, and using two series of F_t estimates, one obtained from demographic parameters, the other from molecular markers. Overall, V_d was slightly less than expected, perhaps as a result of some characters being subject to weak stabilizing selection, but the general picture was one of close concordance with the prediction of the neutral additive model. Plots of phenotypic line means, from the parental generation through to F₃, illustrate that the observed differentiation was essentially entirely due to the initial sampling event. Egg weight showed a similar pattern of differentiation. © 2006 The Linnean Society of London, Biological Journal of the Linnean Society, 2006, 89 , 107–115.     

Variation in the ribosomal RNA genes among individuals of Vicia faba

Summary Length heterogeneity in the ribosomal repeat of Vicia faba is due to the presence of variable numbers of a 325 bp subrepetitive element within the nontranscribed spacer region. The distribution of size classes among 88 individuals within a population was investigated by blot-hybridization. We find that individual plants can exhibit more than 20 size classes and that hybridization patterns are highly diverse from individual to individual, more so than for any species so far investigated. In contrast, no such differences are observed in patterns for different tissues from a single plant or from parental to F₁ generation. Some changes were observed in the F₂ generation. We conclude that unequal recombination can give rise to the diversity that we observe for the V. faba rDNA repeats.