Evolution of developmental homeostasis inDrosophila mojavensis

Summary Variation in life histories among populations of cactophilicDrosophila mojavensis has been hypothesized to be a by-product of a shift to one of two alternate host plants. When cultured on the ancestral and a secondary host cactus, a Baja population expressed shorter development times and smaller thorax sizes than a mainland population, but viability did not differ. Comparisons with all reciprocal F₁ and F₂ crosses between populations revealed that genetic differences in development time and thorax size were largely additive. Homeostasis in these life history traits was population specific, except for viability. Homeostasis in development time was greater in the Baja population than in the other crosses, suggesting dominance for decreased homeostasis in the mainland population. Underdominance in viability homeostasis of the F₁ hybrids suggested some incompatibility between populations. Homeostasis in thorax size was greater in females than in males and differed among parental populations. Maintenance of heritable differences and genetic variation for homeostasis in these traits suggested a role for cactus-specific differences in environmental uncertainty caused by variation in breeding site duration and abundance in nature.     

Epistasis in maize (Zea mays L.)

Summary Three flint and three dent maize (Zea mays L.) inbred lines, their possible F₁ crosses, F₂ and backcross progenies, and all possible three-way crosses were evaluated in a three-year experiment for yield, ear moisture, and plant height. The purpose was to estimate genetic parameters in European breeding materials from (i) generation means analysis, (ii) diallel analysis of generation means, and (iii) analysis of F₁ and three-way cross hybrids. Method (i) was based on the F-metric model and methods (ii) and (iii) on the Eberhart-Gardner (1966) genetic model; both models extended for heterotic maternal effects.Differences among generation means for yield and plant height were mainly attributable to dominance effects. Epistatic effects were significantly different from zero in a few crosses and considerably reduced heterosis in both traits. Additive x additive and domiance x dominance effects for yield were consistently positive and negative, respectively. Significant maternal effects were established to the advantage of generations with a heterozygous seed parent. In the diallel analysis, mean squares for dominance effects were greater than for additive effects for yield and plant height but smaller for ear moisture. Though significant for yield and plant height, epistatic variation was small compared to additive and dominance variation. Estimates of additive x additive epistasis for yield were significantly negative in 11 of 15 crosses, suggesting that advantageous gene combinations in the lines had been disrupted by recombination in the segregating generations. The analysis of hybrids supported the above findings regarding the analysis of variance. However, the estimates of additive x additive epistasis for yield were considerably smaller and only minimally correlated with those from the diallel analysis. Use of noninbred materials as opposed to materials with different levels of inbreeding is considered the main reason for the discrepancies in the results.     

Plant genetic differences influence herbivore community structure: evidence from a hybrid willow system

To determine the influence of plant genetic variation on community structure of insect herbivores, we examined the abundances of 14 herbivore species among six genetic classes of willow: Salix eriocephala, S. sericea, their F₁ and F₂ interspecific hybrids, and backcross hybrids to each parental species. We placed 1-year-old plants, grown from seeds generated from controlled crosses, in a common garden. During the growing season, we censused gall-inducing flies and sawflies, leaf-mining insects, and leaf-folding Lepidoptera to determine the community structure of herbivorous insects on the six genetic classes. Our results provided convincing evidence that the community structure of insect herbivores in this hybrid willow system was shaped by genetic differences among the parental species and the hybrid genetic classes. Using MANOVA, we detected significant differences among genetic classes for both absolute and relative abundance of herbivores. Using canonical discriminant analysis, we found that centroid locations describing community structure of the insect herbivores differed for each genetic class. Moreover, the centroids for the four hybrid classes were located well outside of the range between the centroids for the parental species, suggesting that more than additive genetic effects of the two parental species influenced community formation on hybrid classes. Line-cross analysis suggested that plant genetic factors responsible for structuring the herbivore community involved epistatic effects, as well as additive and dominance effects. We discuss the ramifications of these results in regard to the structure of insect herbivore communities on plants and the implications of our findings for the evolution of interspecific interactions.     

Genetic variation and hybridization in SwedishSchoenus (Cyperaceae)

Schoenus ferrugineus andS. nigricans have restricted distributions in Sweden and are almost exclusively confined to calcareous fen habitats. AtS. nigricans sites,S. ferrugineus is usually also present, and hybrids are frequently found. In this report, I used allozymes to estimate the amount of gene flow between the two species, and to compare the partitioning of genetic diversity in each of them. Thirteen loci were analysed at eight different enzyme systems. Seven loci were variable between or within the species. The two species had completely different alleles at two of the seven variable loci, whereas there was overlap at five loci. In all, 22 different alleles were found. Six of these alleles were confined toS. nigricans, and five alleles were confined toS. ferrugineus. Nei's genetic identity was 0.55.—InS. ferrugineus, three loci (23%) were polymorphic, and the average number of alleles per polymorphic locus was 2.0 (each polymorphic locus had two alleles). InS. nigricans, three loci (23%) were polymorphic, and the average number of alleles per polymorphic locus was 2.3.—The proportion of genetic diversity due to variation among sites (G _ST) was fairly similar in the two species, mean over loci = 0.12 inS. ferrugineus and 0.15 inS. nigricans. However, the proportion of genetic diversity due to variation among individuals within sites (G _IS) differed markedly between the two species, mean over loci = 0.54 inS. ferrugineus and 0.17 inS. nigricans. Accordingly, there was a much higher individual heterozygosity inS. nigricans than inS. ferrugineus. — Most hybrids were interpreted as F₁ hybrids. However, a small proportion, 0.5–1.6 %, were F_n hybrids or back-crosses.—On the Swedish mainland, all former occurrences ofS. nigricans are extinct, but viable hybrids are still present at a few sites in southernmost Sweden.     

QTL analysis of flowering time inArabidopsis thaliana

Quantitative trait loci (QTL) analyses based on restriction fragment length polymorphism maps have been used to resolve the genetic control of flowering time in a cross between twoArabidopsis thaliana ecotypes H51 and Landsbergerecta, differing widely in flowering time. Five quantitative trait loci affecting flowering time were identified in this cross (RLN1-5), four of which are located in regions containing mutations or loci previously identified as conferring a late-flowering phenotype. One of these loci is coincident with theFRI locus identified as the major determinant for late flowering and vernalization responsiveness in theArabidopsis ecotype Stockholm.RLN5, which maps to the lower half of chromosome five (between markers mi69 and m233), only affected flowering time significantly under short day conditions following a vernalization period. The late-flowering phenotype of H51 compared to Landsbergerecta was due to alleles conferring late flowering at only two of the five loci. At the three other loci, H51 possessed alleles conferring early flowering in comparison to those of Landsbergerecta. Combinations of alleles conferring early and late flowering from both parents accounted for the transgressive segregation of flowering time observed within the F₂ population. Three QTL,RLN1,RLN2 andRLN3 displayed significant genotype-by-environment interactions for flowering time. A significant interaction between alleles atRLN3 andRLN4 was detected.     

Genes for dwarfing and photoperiod flowering response inPharbitis nil choisy

Dwarfing and sensitivity to the duration of a single inductive dark period for flowering ofPharbitis nil in F₂ progeny of a cross between the tall strain Tendan, and the dwarf, Kidachi appear to be controlled by the alleles at two independent loci. Progeny of a similar cross between the tall strain Violet and the dwarf Kidachi at F₂ and F₃ also showed single locus segregation for tall: dwarf plants. In this cross, differences in photoperiodic response could be identified in F₃ families but they were not simply inherited. There was some evidence of difficulties with classification of the F₂ plants, but also, the flowering of the F₁ between the two less sensitive strains Tendan and Violet indicated complex inheritance of their photoperiodic response. Complementary dominant alleles at three independent loci may be necessary for flowering in even shorter dark periods with the sensitive strain Kidachi. The dwarf strain Kidachi has a reduced gibberellin (GA) content (Barendse and Lang 1972), it flowers in a short dark period without terminal flowering, and it responds positively to GA application both for flowering and growth. However, since control of dwarfing and photoperiodic sensitivity can be separated genetically, there is no strick link between the gibberellin responsiveness of Kidachi for its growth and flowering. Despite the complexity of flowering genetics in Violet×Kidachi, a short-dark-period-sensitive, terminal flowering and tall F₇ line was obtained in a pedigree previously held heterozygous for the dwarf: tall character but not selected for flowering time. Thus, flowering in a short dark period can also be obtained in the presence of the non-dwarfing allele from strain Violet, again demonstrating genetic independence.     

Mutationsversuche an Kulturpflanzen

Summary The genetic behaviour of some X-ray induced early heading mutants of both spring and winter barley was studied. In crosses with original varieties all mutants showed monogenic segregation ratios in F₂-generations. The early heading date of four mutants was found to be due to a dominant allele, while six mutants are dependent on recessive alleles.In crosses of some mutants one with another four different loci for early heading were identified. For them the genetic symbolmat (matura) resp.Mat is proposed.The combination-effect of two non-allelic genes is always additive and manifests itself in a marked transgression. Plants with such genotypes seem to be useless for breeding purposes.

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GENETIC ARCHITECTURE OF ISOLATION BETWEEN TWO SPECIES OF SILENE WITH SEX CHROMOSOMES AND HALDANE'S RULE

Examination of the genetic architecture of hybrid breakdown can provide insight into the genetic mechanisms of commonly observed isolating phenomena such as Haldane's rule. We used line‐cross analysis to dissect the genetic architecture of divergence between two plant species that exhibit Haldane's rule for male sterility and rarity, Silene latifolia and Silene diclinis. We made 15 types of crosses, including reciprocal F₁, F₂, backcrosses, and later‐generation crosses, grew the seeds to flowering, and measured the number of viable ovules, proportion of viable pollen, and sex ratio. Typically, Haldane's rule for male rarity in XY animal hybrids is explained by interactions involving recessive X‐linked alleles that are deleterious when hemizygous (dominance theory), whereas sterility is explained by rapid evolution of spermatogenesis genes (faster‐male evolution). In contrast, we found that the genetic mechanisms underlying Haldane's rule between the two Silene species did not follow these conventions. Dominance theory was sufficient to explain male sterility, but male rarity likely involved faster‐male evolution. We also found an effect of the neo‐sex chromosomes of S. diclinis on the extreme rarity of some hybrid males. Our findings suggest that the genetic architecture of Haldane's rule in dioecious plants may differ from those commonly found in animals.     

Pattern of genetic differentiation in two Isophya species (Orthoptera: Tettigonoidea) in north-east Hungary

Allozyme polymorphism was studied in two populations of Isophya kraussi and Isophya stysi. Both species are flightless and have low dispersal ability. As a consequence, we expected high level of genetic differentiation among their local populations. Samples were collected in three regions of Hungary. Enzyme polymorphism was investigated at 10 loci (Aldox, Est, Got, Gpdh, Hk, Idh, Mdh, Me, Pgi and Pgm) in both species. High levels of polymorphism were detected in all samples. Gpdh proved to be diagnostic as there were no common alleles in the two species. At four further loci (Got, Hk, Mdh and Me), the two species had one common allele together with one or more differentiating alleles. We detected high F _IT values implying a high level of genetic variation. The positive F _IS values suggested a tendency of heterozygote deficiency in both species. The highly significant overall F _ST values indicated clear genetic differentiation among the local populations. Thus our results confirmed the taxonomic status of these two species. The dendrogram constructed on the basis of Nei's genetic distances and the results of the PCA analyses fully confirmed those obtained by F-statistics.     

Genetic analysis of Verticillium wilt tolerance in cotton using pedigree data from three crosses

Summary Three crosses and descendant generations were used in a field study of the inheritance of tolerance to Verticillium wilt, caused by Verticillium dahliae Kleb., in upland cotton (Gossypium hirsutum L.). The tolerant cultivar Acala SJC-1 was crossed to more susceptible parents, breeding line S5971 and cultivars Acala 4-42 and Deltapine 70. Seven generations were evaluated for each cross: the two parents (P₁ and P₂), F₁; F₂, F₃, and reciprocal backcrosses (B₁ and B₂). The genetic control of tolerance in these crosses appears to involve more than one gene, based on an unsatisfactory fit to expected phenotypic distributions for the generations under a single-locus model. An analysis of generation means indicated that pooled additive and pooled dominance effects over loci were adequate to explain the variation among generations for crosses of SJC-1 X S5971 and SJC-1 X DPL70. Tolerance in these crosses appeared to be controlled by recessive factors. For the SJC-1 X 4-42 cross, an adequate fit to a digenic epistatic model was not possible, and none of the genetic parameters except the F₂ mean were significant. Heritabilities for tolerance to Verticillium wilt, determined from regressions of F₃ progeny on F₂ parents for the crosses of SJC-1 X S5971 and SJC-1 X DPL70, ranged from 0.12 to 0.28. Therefore, individual plant selection for improved tolerance is expected to be inefficient.Contribution from the Department of Botany and Plant Sciences, University of California, Riverside, CA 92521, USA, to be included in a dissertation by the senior author in partial fulfillment of the Ph. D. degree     

Validation of the <Emphasis Type="Italic">VRN-H2</Emphasis>/<Emphasis Type="Italic">VRN-H1</Emphasis> epistatic model in barley reveals that intron length variation in <Emphasis Type="Italic">VRN-H1</Emphasis> may account for a continuum of vernalization sensitivity

The epistatic interaction of alleles at the VRN-H1 and VRN-H2 loci determines vernalization sensitivity in barley. To validate the current molecular model for the two-locus epistasis, we crossed homozygous vernalization-insensitive plants harboring a predicted “winter type” allele at either VRN-H1 (Dicktoo) or VRN-H2 (Oregon Wolfe Barley Dominant), or at both VRN-H (Calicuchima-sib) loci and measured the flowering time of unvernalized F₂ progeny under long-day photoperiod. We assessed whether the spring growth habit of Calicuchima-sib is an exception to the two-locus epistatic model or contains novel “spring” alleles at VRN-H1 (HvBM5A) and/or VRN-H2 (ZCCT-H) by determining allele sequence variants at these loci and their effects relative to growth habit. We found that (a) progeny with predicted “winter type” alleles at both VRN-H1 and VRN-H2 alleles exhibited an extremely delayed flowering (i.e. vernalization-sensitive) phenotype in two out of the three F₂ populations, (b) sequence flanking the vernalization critical region of HvBM5A intron 1 likely influences degree of vernalization sensitivity, (c) a winter habit is retained when ZCCT-Ha has been deleted, and (d) the ZCCT-H genes have higher levels of allelic polymorphism than other winterhardiness regulatory genes. Our results validate the model explaining the epistatic interaction of VRN-H2 and VRN-H1 under long-day conditions, demonstrate recovery of vernalization-sensitive progeny from crosses of vernalization-insensitive genotypes, show that intron length variation in VRN-H1 may account for a continuum of vernalization sensitivity, and provide molecular markers that are accurate predictors of “winter vs spring type” alleles at the VRN-H loci.     

Use of single-primer DNA amplifications in genetic studies of peanut (Arachis hypogaea L.)

A recent approach to detecting genetic polymorphism involves the amplification of genomic DNA using single primers of arbitrary sequence. When separated electrophoretically in agarose gels, the amplification products give banding patterns that can be scored for genetic variation. The objective of this research was to apply these techniques to cultivated peanut (Arachis hypogaea L.) and related wild species to determine whether such an approach would be feasible for the construction of a genetic linkage map in peanut or for systematic studies of the genus. Two peanut cultivars, 25 unadapted germplasm lines of A. hypogaea, the wild allotetraploid progenitor of cultivated peanut (A. monticola), A. glabrata (a tetraploid species from section Rhizomatosae), and 29 diploid wild species of Arachis were evaluated for variability using primers of arbitrary sequence to amplify segments of genomic DNA. No variation in banding pattern was observed among the cultivars and germplasm lines of A. hypogaea, whereas the wild Arachis species were uniquely identified with most primers tested. Bands were scored (+/–) in the wild species and the PAUP computer program for phylogenetic analysis and the HyperRFLP program for genetic distance analysis were used to generate dendrograms showing genetic relationships among the diploid Arachis species evaluated. The two analyses produced nearly identical dendrograms of species relationships. In addition, approximately 100 F₂ progeny from each of two interspecific crosses were evaluated for segregation of banding patterns. Although normal segregation was observed among the F₂ progeny from both crosses, banding patterns were quite complex and undesirable for use in genetic mapping. The dominant behavior of the markers prevented the differentiation of heterozygotes from homozygotes with certainty, limiting the usefulness of arbitrary primer amplification products as markers in the construction of a genetic linkage map in peanut.     

Mapping loci controlling vernalization requirement in Brassica rapa

Brassica cultivars are classified as biennial or annual based on their requirement for a period of cold treatment (vernalization) to induce flowering. Genes controlling the vernalization requirement were identified in a Brassica rapa F₂ population derived from a cross between an annual and a biennial oilseed cultivar by using an RFLP linkage map and quantitative trait locus (QTL) analysis of flowering time in F₃ lines. Two genomic regions were strongly associated with variation for flowering time of unvernalized plants and alleles from the biennial parent in these regions delayed flowering. These QTLs had no significant effect on flowering time after plants were vernalized for 6 weeks, suggesting that they control flowering time through the requirement for vernalization. The two B. rapa linkage groups containing these QTLs had RFLP loci in common with two B. napus linkage groups that were shown previously to contain QTLs for flowering time. An RFLP locus detected by the cold-induced gene COR6.6 cloned from Arabidopsis thaliana mapped very near to one of the B. rapa QTLs for flowering time.     

Genetic studies of yield contributing traits in Amaranthus

Summary Genetical studies on grain yield and its contributing traits were made in a six parent complete diallel in the F₁ and F₂ generations of one of the most widely grown grain species of grain amaranths (Amaranthus hypochondriacus L.). Graphical analysis indicated that epistasis exists for 1,000-grain weight in the F₁. Grain weight/panicle, yield/plant and harvest index indicated absence of non-allelic gene interaction. The harvest index in the F₁ and F₂ and grain weight/ panicle, 1,000-grain weight, yield/plant in the F₂ appeared to be controlled by overdominance effects. Higher grain yield appeared to be associated with dominant genes. Both additive and non-additive gene effects were responsible for the genetic variation in the diallel population. However, dominance variance was more important than additive variance in grain yield/ plant and harvest index in the F₁ and F₂. For 1,000-grain weight additive genetic variance was more important in the F₁ and non-additive in F₂. There was overdominance of a consistent nature in the two analyses for harvest index in the F₁ and F₂, grain weight/ panicle, 1,000-grain weight and yield/plant in the F₂ and partial dominance for 1,000-grain weight in the F₁.     

The effect of genome and sex on recombination rates in Pennisetum species

The effects of homoeology and sex on recombination frequency were studied in crosses between cultivated pearl millet, Pennisetum glaucum, and two wild subspecies, P. violaceum and P. mollissimum. For the two wild x cultivated crosses, reciprocal three-way crosses were made between the F₁ hybrid and an inbred line (Tift 23DB₁). The three-way cross populations were mapped to produce a female map of each wide cross (where the F₁ was the female) and a male map (where the F₁ was the male). Total genetic map lengths of the two inter-subspecies crosses were broadly similar and around 85 % of a comparable intervarietal map. In the P. glaucumxP. mollissimum crosses, the map was further shortened by a large (40 cM) inversion in linkage group 1. Comparison of the recovered recombinants from male and female meiocytes showed an overall trend for the genetic maps to be longer in the male (10%) in both inter-subspecific crosses; however, analysis of individual linkage intervals showed no significant differences. Gametophytic selection was prevalent, and sometimes extreme, for example 121 in favour of wild alleles in the P. glaucumxP. mollissimum male recombinant population. One of the loci which determines panicle type in cultivated pearl millet and wild relatives, H, was mapped 9 cM from Xpsm812 on linkage group 7 in the P. violaceum cross.     

Allozyme variation in the sympatric ferns Culcita macrocarpa and Woodwardia radicans at the northern extreme of their ranges

Isozymes were used to study genetic variation in the clonal ferns Culcita macrocarpa and Woodwardia radicans in the northwestern Iberian Peninsula, their northern distributional limit. Despite their high chromosome numbers, both species were isozymic diploids. In C. macrocarpa all 18 resolved loci were monomorphic, with the same allele in all ramets from all populations. In W. radicans only two of the 16 interpreted loci were polymorphic, with two alleles per locus; ramet-level genotypes showed Hardy-Weinberg equilibrium, indicating an intergametophytic mating system; the number of genets distinguished was 1–3 per population; and among-population variation was low (F _ST = 0.231), suggesting effective gene flow (i.e. spore exchange). More generally, the very low (W. radicans, H _T = 0.012) or zero (C. macrocarpa) genetic diversity detected in the present study may be due to genetic drift associated with the reduction of populations in the last glaciation, and to founder effects in the subsequent Holocene expansion.     

Gene effects for fibre properties in upland cotton (Gossypium hirsutum L.)

Summary Six populations — P₁,P₂,F₁,F₂,B₁ and B₂ — each of five Upland Cotton (Gossypium hirsutum L.) crosses were used to evaluate gene effects in the inheritance of fibre properties by Gamble's six-parameter model for the analysis of generation means. Partial dominance of long fibres over short fibres and of mature fibres over immature fibres was found in the material studied. Overdominance in gene action governed fibre fineness while additive gene action governed the fibre strength. Besides additive and dominance effects, significant epistasis was present in all crosses. These results indicate a significant potential for improving fibre properties through reciprocal recurrent selection.     

Segregation analysis and RFLP mapping of the R1 and R3 alleles conferring race-specific resistance to Phytophthora infestans in progeny of dihaploid potato parents

Phytophthora infestans (Mont.) de Bary is the most important fungal pathogen of the potato (Solanum tuberosum). The introduction of major genes for resistance from the wild species S. demissum into potato cultivars is the earliest example of breeding for resistance using wild germplasm in this crop. Eleven resistance alleles (R genes) are known, differing in the recognition of corresponding avirulence alleles of the fungus. The number of R loci, their positions on the genetic map and the allelic relationships between different R variants are not known, except that the R1 locus has been mapped to potato chromosome V The objective of this work was the further genetic analysis of different R alleles in potato. Tetraploid potato cultivars carrying R alleles were reduced to the diploid level by inducing haploid parthenogenetic development of 2n female gametes. Of the 157 isolated primary dihaploids, 7 set seeds and carried the resistance alleles R1, R3 and R10 either individually or in combinations. Independent segregation of the dominant R1 and R3 alleles was demonstrated in two F₁ populations of crosses among a dihaploid clone carrying R1 plus R3 and susceptible pollinators. Distorted segregation in favour of susceptibility was found for the R3 allele in 15 of 18 F₁ populations analysed, whereas the RI allele segregated with a 1:1 ratio as expected in five F₁ populations. The mode of inheritance of the R10 allele could not be deduced as only very few F₁ hybrids bearing R10 were obtained. Linkage analysis in two F₁ populations between R1, R3 and RFLP markers of known position on the potato RFLP maps confirmed the position of the R1 locus on chromosome V and localized the second locus, R3, to a distal position on chromdsome XI.     

Genes for Dwarfness in Wheat, TRITICUM AESTIVUM L

The genetic control of plant height was studied in crosses of four spring wheats involving the standard height variety Ramona 50 and short-statured selections Olesen, D6301, and D6899. Data from parent, F₁, F₂, and F₃ populations indicated that four independently segregating loci account for most of the differences among the four varieties. Two major genes of a highly recessive nature condition reduced height in Olesen and the Norin 10 derivative D6301. Olesen also carries a third dwarfing gene which is partially dominant in its effects over genes for tallness. This gene, or a gene that acts in a similar manner, is also present in the standard height variety Ramona 50. Dwarfing in D6899, a derivative of Tom Thumb, is controlled primarily by a single gene with mainly additive effects which is not present in any of the other three varieties.

Genetic components estimated from generation means (parental, F₁, F₂, F₃, and backcross) indicated that additive gene effects were the major component of variation in four of the six crosses, and of similar magnitude to dominance effects in another cross. The primary source of genetic variation in the cross Olesen x D6899 was due to epistasis with both additive x additive and dominance x dominance effects of major importance. The results of the generation mean analyses were consistent with the models for major-gene control of plant height based on segregation patterns.