Genetic parameters for agronomic characteristics. I. Early and intermediate breeding populations of true potato seed

The original variation in the source population as well as the selection method may influence the genetic variation in further cycles of genetic improvement. Therefore, the objectives of this research were to determine genetic parameters (variance components and heritability) in source and intermediate stages of a true potato seed (TPS) breeding population and to calculate the genetic and phenotypic correlations in this breeding material developed by the Centro Internacional de la Papa (CIP). The intermediate stage was derived from a source population adapted to the warm lowland tropics plus introduction of exotic germplasm from North America and Europe. Non-additive genetic variation was almost nil for plant survival, tuber yield and tuber shape uniformity in both stages of the breeding population and no quantitative genetic variation for uniformity of tuber color was observed in both source and intermediate breeding materials. Heritability was higher in the intermediate stage than in the source population for plant survival (0.86 vs 0.66), tuber yield (0.30 vs 0.14) and tuber shape (0.77 vs 0.51), but it was the reverse for tuber uniformity (0.11 vs 0.72). These results suggest that potato breeders at CIP were able to keep enough genetic variation for most important characteristics for potato production from true seed in their intermediate breeding materials by adding new sources of variation to the original breeding population. Additive genetic and phenotypic correlations were significant and positive between plant vigor after transplanting and tuber yield, and tuber shape and tuber uniformity, which suggest that high yielding offspring result from early vigorous growth, and that tuber uniformity could depend on tuber shape uniformity in this breeding material.     

Early generation selection for agronomic characters in a potato breeding programme

Summary A random sample of seedlings representing high, medium and poor vigour was studied for tuber colour, tuber shape, eye depth, tuber cracking, tuber yield per plant, average tuber weight and number of tubers per plant in four successive generations (F₁, F₁, F₁C₂, and F₁C₃). Based on the performance of vigour groups in various generations and inter-generation correlation coefficients, we propose a procedure for the elimination of unproductive genotypes early in the breeding programme. The data indicates that seedlings of poor vigour can be discarded at the seedling stage prior to transplantation in the field. The rejection of clones on the basis of tuber colour, tuber shape, eye depth and tuber cracking can also be initiated at the seedling stage. For tuber yield and average tuber weight a negative selection (rejection of poor phenotypes) is suggested from the first clonal generation and for number of tubers, from second clonal generation, until statistically sound replicated trials can be conducted for carrying out positive selection.     

Progeny selection for agronomic characters in early generations of a potato breeding programme

 Repeatabilities of progeny means, and the univariate cross prediction method were used to study the effectiveness of progeny selection for agronomically important characters in early generations of potato (Solanum tuberosum L.) breeding. The study was based on 90 progenies (72 crosses+18 selfs) evaluated for three successive generations, i.e. seedling, first clonal and second clonal generations. Repeatabilities of progeny means were measured as correlation coefficients between generations. In the univariate cross prediction method, progeny means and within-progeny standard deviations were used to calculate the proportions of clones exceeding the target values, and correlation coefficients between generations for predicted and observed proportions of clones, were calculated. Population means varied from generation to generation. Correlation coefficients between generations for progeny means for most of the characters were significant, but moderate. These were higher than the correlation coefficients between predicted and observed proportions of clones exceeding the target values. The possibility of using progeny means as a selection parameter to reduce the number of genotypes to be examined in later stages by rejecting the poor crosses in seedling generation is discussed. Received: 8 January 1997/Accepted: 28 February 1997     

Inbreeding and true seed in tetrasomic potato. IV. Synthetic cultivars

A true potato seed (TPS) synthetic cultivar from open pollination may broaden the genetic base in potato. A synthetic cultivar can also reduce the effect of inbreeding and maintain the productivity from generation to generation. Selected tetraploid parental genotypes, based on combining-ability tests, were chosen for isolated polycrosses to obtain tetraploid offspring. A series of experiments were undertaken to investigate the development of these TPS synthetic populations in two Peruvian locations: San Ramon, a rainfed, humid, mid-altitude environment, and La Molina, an arid, coastal environment under irrigation. Natural open-pollinated synthetics or control-mixture synthetics, involving two to six parents, had a similar tuber set but plant survival and tuber weight were higher in control synthetics including two or six parents. The results suggest that two-parent open-pollinated TPS synthetics may be a feasible option by selecting the right parents for the base population. A subsequent experiment showed that four-parent control synthetics (i.e. double crosses) were the best for tuber weight and size, followed by the two-parent control synthetics. Synthetic breeding populations could be shared with other breeders, who in cooperation with their local farmers may select promising genotypes for further cultivar release. Received: 15 December 2000 / Accepted: 4 April 2001     

Inbreeding and true seed in tetrasomic potato. II. Selfing and sib-mating in heterogeneous hybrid populations of Solanum tuberosum

 Inbreeding depression may affect the performance of consecutive generations of potatoes propagated by true potato seed (TPS). The effect of inbreeding was established using selfed and sib-mated generations of five TPS families. Correlation coefficients were calculated between the level of inbreeding and different traits. Inbreeding depression was expressed mainly by pollen viability, as measured by its stainability (r=−0.912, P<0.01), and tuber yield (r=−0.837, 0.01<P<0.05). The results also indicated that without unavoidable selection inbreeding depression is expected to be more evident. Furthermore, the TPS families responded quite differently to inbreeding depression. They did not show the same amount of depression for yield as they did for the characters concerning fertility. The high tuber-yielding families displayed a greater inbreeding depression for tuber yield than the lower-yielding families. Received: 10 November 1997 / Accepted: 3 March 1998     

Producing commercially attractive,uniform true potato seed progenies: the influence of breeding scheme and parental genotype

The production of attractive, uniform true potato seed (TPS) progenies was investigated. Four breeding schemes were compared: intercrossing tetraploid cultivars (cv x cv); doubled dihaploids x cultivars (ddh x cv); cultivars x diploid unreduced-gamete producers (cv x FDR) and doubled dihaploids x diploid unreduced-gamete producers (ddh x FDR). Fifty three progenies and five clones were grown in a glasshouse in a randomised complete block design with three replicates of 25 plants per progeny and clone. Each plant's tubers were counted and the colour, shape, quality of skin finish, flesh colour, and commencai attractiveness (which includes yield) recorded. The most uniform progenies were also selected by visual comparison with the clones. For mean attractiveness, differences (P < 0.001) between breeding schemes and between progenies within breeding schemes were detected. The cv x cv and cv x FDR progenies were more attractive than clonal controls. There were significant additive and non-additive effects for attractiveness in all breeding schemes except cv x FDR. There were between-progeny differences (P < 0.001) for uniformity for all characters. Progenies uniform for one character could be variable for other traits. Breeding schemes gave different levels of uniformity (P < 0.001) for all characters except shape and flesh colour, but none gave low levels of variation for all traits. Doubled-dihaploid parents increased the variation in progenies. There were uniformity differences (P < 0.001) between progenies within breeding schemes for all characters. Evidence of additive and nonadditive genetic variation for uniformity in all traits was detected. In each breeding scheme, parents with good general combining ability (GCA) for uniformity in several characters were identified. Visually selected uniform progenies had parents with good GCAs for uniformity in a range of traits and high specific combining abilities (SCAs) for several traits. A desynaptic first-division restitution (FDR) clone and a male-sterile doubled-dihaploid clone had the best GCAs for tuber uniformity in TPS progenies. Achieving multitrait uniformity in TPS is problematic but may be aided by the selection of parents with GCAs for uniformity coupled with progeny testing to allow for non-additive effects.     

Inbreeding and true seed in tetrasomic potato. III. Early selection for seedling vigor in open-pollinated populations

Tuber yield in open-pollinated (OP) offspring of potato cultivars appears to be correlated with the proportion of hybrids within each population. OP true potato seed (TPS) from ten selected clones was tested in an experiment at two Peruvian locations, San Ramón (mid-altitude warm tropics) and Huancayo (highland cool tropics), to investigate the early selection effect on OP populations. Another experiment in La Molina (dry coast) was included to confirm the results from these other two locations. The results clearly showed that plant survival, tuber yield and berry number can be increased by a strong early selection. Likewise, the results suggested that the most-promising OP offspring for TPS production could be derived from tetraploid clones with a high rate of outcrossing. This experiment demonstrates that high tuber yield may be obtained in OP TPS cultivars with little effort, using early selection for seedling vigor in the nursery. Genetic interpretation of this response to early selection indicates that both the intensity of selection and non-additive genetic variation for tuber yield account for these observed gains. Received: 15 December 2000 / Accepted: 4 April 2001     

Discovery of a synaptic mutant in potato haploids and its usefulness for potato breeding

Summary A synaptic mutant was found in haploids (2n=2x=24) extracted from the Mexican potato variety Atzimba (2n=4x=48). The mutant is inherited as a simple Mendelian recessive, designated sy4. Meiotic abnormalities of the mutant during microsporogenesis include: poor synapsis at pachytene; high frequency of univalents at diakinesis; elongated and curved spindles and univalents being scattered over the spindles at metaphase I and anaphase I; abnormal chromosome distribution at anaphase I; and production of sterile pollen, presumably due to unbalanced chromosome complement. The expression of sy4 in megasporogenesis was also detected. The sy4 mutant is very useful for potato breeding when combined with another meiotic mutant, parallel spindles (ps), because haploids homozygous for sy4 and ps produce fertile 2n pollen which transmit almost intact genotypes of the parents to the progenies. Thus, the meiotic mutants provide a powerful breeding method for maximizing heterozygosity and epistasis. They can also provide a very efficient method of transferring diploid germplasm, which has desired characteristics efficiently combined at the 2x level, to tetraploids. Many haploids have been identified with 2n pollen production by ps alone or by sy4 and ps, vigorous growth and good flowering, and a high level of resistance to late blight. The importance of a further search for meiotic mutants and their use for breeding is discussed.     

Genetic diversity in a seed production population vs. natural populations of Sitka Spruce

Isozyme analysis was applied to estimate the level of variation and the genetic structure of a seed-production population (i.e., seed orchard) and 10 range-wide natural populations of Sitka spruce (Picea sitchensis (Bong.) Carr.). Gene diversity and heterozygosity estimates were comparatively high in both the seed orchard and the natural populations studied. The seed orchard population showed a significantly higher number of alleles per locus and percentage of polymorphic loci. Though not significant, mean heterozygosity of the seed orchard was higher than that observed for all natural populations. Genetic distance analysis indicated that the seed-orchard population was genetically similar to three natural populations from which the parent trees were selected. Parent trees sampling breadth has been identified as the major cause for the observed increased level. The impact of recurrent selection and seed orchard biology and management on maintaining the genetic diversity is discussed.     

Inbreeding and true seed in tetrasomic potato. I. Selfing and open pollination in Andean landraces (Solanum tuberosum Gp. Andigena)

 True potato seed (TPS) may be an alternative method of potato production in developing countries. A breeding method for the sexual propagation of this vegetatively propagated crop should consider the development of parental lines and the type of cultivar to be released. Open-pollinated (OP) cultivars seem to be an inexpensive procedure to produce potato from true seed. However, OP progenies are the result of selfing and outcrossing in male-fertile tetraploid potatoes. The aim of the present research was to establish the effect of inbreeding and open pollination in TPS. Ten Andigena clones were used as parental material to derive hybrid (S₀), inbred (S₁ and S₂), and open-pollinated (OP₁ and OP₂) generations. Significant differences among generations were found for pollen production, pollen viability (as determined by its stainability with aceto-carmine glycerol), number of flowers and berries plant^-1, number of seeds berry^-1, weight of 1000 seeds, and tuber yield plant^-1. The parental populations were significantly different for most of the traits, but not for flower production and berry weight. The interaction of population ×generation was significant for pollen and seed production as well as for weight for 1000 seeds. All the traits evaluated except seed weight showed a strong inbreeding depression, while the OP progenies had intermediate values between the S₀ and the S₁. This demonstrates that open pollination in potatoes is not exclusively the product of selfing; it also results from outcrossing. Received: 10 November 1997 / Accepted: 3 March 1998     

Genetic parameters for growth, reproductive and maternal traits in a multibreed meat sheep population

The genetic parameters for growth, reproductive and maternal traits in a multibreed meat sheep population were estimated by applying the Average Information Restricted Maximum Likelihood method to an animal model. Data from a flock supported by the Programa de Melhoramento Genético de Caprinos e Ovinos de Corte (GENECOC) were used. The traits studied included birth weight (BW), weaning weight (WW), slaughter weight (SW), yearling weight (YW), weight gain from birth to weaning (GBW), weight gain from weaning to slaughter (GWS), weight gain from weaning to yearling (GWY), age at first lambing (AFL), lambing interval (LI), gestation length (GL), lambing date (LD - number of days between the start of breeding season and lambing), litter weight at birth (LWB) and litter weight at weaning (LWW). The direct heritabilities were 0.35, 0.81, 0.65, 0.49, 0.20, 0.15 and 0.39 for BW, WW, SW, YW, GBW, GWS and GWY, respectively, and 0.04, 0.06, 0.10, 0.05, 0.15 and 0.11 for AFL, LI, GL, LD, LWB and LWW, respectively. Positive genetic correlations were observed among body weights. In contrast, there was a negative genetic correlation between GBW and GWS (-0.49) and GBW and GWY (-0.56). Positive genetic correlations were observed between AFL and LI, LI and GL, and LWB and LWW. These results indicate a strong maternal influence in this herd and the presence of sufficient genetic variation to allow mass selection for growth traits. Additive effects were of little importance for reproductive traits, and other strategies are necessary to improve the performance of these animals.     

AB-QTL analysis in winter wheat: II. Genetic analysis of seedling and field resistance against leaf rust in a wheat advanced backcross population

The present study aimed to localize exotic quantitative trait locus (QTL) alleles for the improvement of leaf rust (P. triticina) resistance in an advanced backcross (AB) population, B22, which is derived from a cross between the winter wheat cultivar Batis (Triticum aestivum) and the synthetic wheat accession Syn022L. The latter was developed from hybridization of T. turgidum ssp. dicoccoides and T. tauschii. Altogether, 250 BC₂F₃ lines of B22 were assessed for seedling resistance against the leaf rust isolate 77WxR under controlled conditions. In addition, field resistance against leaf rust was evaluated by assessing symptom severity under natural infestation across multiple environments. Simultaneously, population B22 was genotyped with a total of 97 SSR markers, distributed over the wheat A, B and D genomes. The phenotype and genotype data were subjected to QTL analysis by applying a 3-factorial mixed model analysis of variance including the marker genotype as a fixed effect and the environments, the lines and the marker by environment interactions as random effects. The QTL analysis revealed six putative QTLs for seedling resistance and seven for field resistance. For seedling resistance, the effects of exotic QTL alleles improved resistance at all detected loci. The maximum decrease of disease symptoms (−46.3%) was associated with marker locus Xbarc149 on chromosome 1D. For field resistance, two loci had stable main effects across environments and five loci exhibited marker by environment interaction effects. The strongest effects were detected at marker locus Xbarc149 on chromosome 1D, at which the exotic allele decreased seedling symptoms by 46.3% and field symptoms by 43.6%, respectively. Some of the detected QTLs co-localized with known resistance genes, while others appear to be as novel resistance loci. Our findings indicate, that the exotic wheat accession Syn022L may be useful for the improvement of leaf rust resistance in cultivated wheat.     

Analysis of QTLs for yield components, agronomic traits, and disease resistance in an advanced backcross population of spring barley.

Hordeum vulgare subsp. spontaneum, the wild progenitor of barley, is a potential source of useful genetic variation for barley breeding programs. The objective of this study was to map quantitative trait loci (QTLs) in an advanced backcross population of barley. A total of 207 BC3 lines were developed using the 2-rowed German spring cultivar Hordeum vulgare subsp. vulgare 'Brenda' as a recurrent parent and the H. vulgare subsp. spontaneum accession HS584 as a donor parent. The lines were genotyped by 108 simple-sequence repeat (SSR) markers and evaluated in field tests for the measurement of grain yield and its components, such as ear length, spikelet number per spike, grain number per spike, spike number, and 1000-grain mass, as well as heading date and plant height. A total of 100 QTLs were detected. Ten QTLs with increasing effects were found for ear length, spikelet number, and grain number per spike. Three QTLs contributed by HS584 were found to significantly decrease days to heading across all years at 2 locations. In addition, 2 QTLs from HS584 on chromosomes 2H and 3H were associated with resistance to leaf rust. Based on genotypic data obtained from this population, 55 introgression lines carrying 1 or 2 donor segments were selected to develop a set of doubled-haploid lines, which will be used to reconfirm and investigate the effects of 100 QTLs for future genetic studies.     

Genetic analyses of agronomic and seed quality traits of synthetic oilseedBrassica napus produced from interspecific hybridization ofB. campestris andB. oleracea

The heritability, the number of segregating genes and the type of gene interaction of nine agronomic traits were analysed based on F₂ populations of synthetic oilseedBrassica napus produced from interspecific hybridization ofB. campestris andB. oleracea through ovary culture. The nine traits—plant height, stem width, number of branches, length of main raceme, number of pods per plant, number of seeds per pod, length of pod, seed weight per plant and 1000-seed weight—had heritabilities of 0.927, 0.215, 0.172, 0.381, 0.360, 0.972, 0.952, 0.516 and 0.987 respectively, while the mean numbers of controlling genes for these characters were 7.4, 10.4, 9.9, 12.9, 11.5, 21.7, 20.5, 19.8 and 6.4 respectively. According to estimated coefficients of skewness and kurtosis of the traits tested, no significant gene interaction was found for plant height, stem width, number of branches, length of main raceme, number of seeds per pod and 1000-seed weight. Seed yield per plant is an important target for oilseed production. In partial correlation analysis, number of pods per plant, number of seeds per pod and 1000-seed weight were positively correlated with seed yield per plant. On the other hand, length of pod was negatively correlated (r = -0.69) with seed yield per plant. Other agronomic characters had no significant correlation to seed yield per plant. In this experiment, the linear regressions of seed yield per plant and other agronomic traits were also analysed. The linear regression equation wasy = 0.074x₈ + 1.819x₉ + 6.72x₁₂ -60.78 (R ² = 0.993), wherex ₈, x₉ and x₁₂ represent number of pods per plant, number of seeds per pod and 1000-seed weight respectively. The experiment also showed that erucic acid and oil contents of seeds from F₂ plants were lower than those of their maternal parents. However, glucosinolate content was higher than that of the maternal plants. As for protein content, similar results were found in the F₂ plants and their maternal parents. It was shown that the four quality traits, i.e. erucic acid, glucosinolate, oil content, and protein content, had heritability values of 0.614, 0.405, 0.153 and 0.680 respectively.     

Genetic dissection of seed shattering,agronomic, and color traits in American wildrice ( Zizania palustris var. interior L.) with a comparative map

A comparative map of American wildrice ( Zizania palustris var. interior L.) was used to identify loci controlling seed shattering, plant height, maturity, tiller number, plant habit, panicle length seed length, and color traits. Two to six significant quantitative-trait-loci (QTLs, P < 0.05) were detected for each trait evaluated, representing the first trait-mapping in wildrice. The chosen population was designed to emphasize the mapping of loci controlling the shattering trait, which is the most important trait in the management of this newly domesticated species. Three loci were detected that controlled the discretely categorized variation between shattering and non-shattering plants. Seed-shattering loci were detected and validated among the F(2) and F(3) generations. A multiple regression model with these three loci described 49.6% of the additive genetic variation. A genetic model with the same three loci including dominance and locus interactions predicted the shattering versus non-shattering phenotype at a success rate of 87%. The comparative map was based on mapped RFLP markers used in white rice ( Oryza sativa L.) and other grass species. Anchor loci provided a reference point for the identification of potential orthologous genes on the basis of white rice mutant loci and consensus grass species QTLs. Candidate orthologous loci were identified among all traits evaluated. The study underscores the benefits of extending trait analysis through comparative mapping, as well as challenges of QTL analysis in a newly domesticated species.     

Genetic parameters in a rubber tree population: heritabilities,genotype-by-environment interactions and multi-trait correlations

Rubber tree breeding programs are mainly driven by selection of individuals with high yield and quality of rubber. Data from 51 open-pollinated progenies tested on six sites in Brazil were analyzed over several traits to estimate the following: genetic parameters such as narrow-sense heritability and additive genetic variance in single- and multi-site analyses, type B correlations to determine the relevance of genotype-by-environment interactions and its effects on alternative selection strategies, additive genetic repeatability correlation for rubber yield based on three consecutive yearly measurements, and type A correlations to evaluate trait-to-trait genetic associations for all measured traits. Average rubber yield (RYm) showed an estimated narrow-sense heritability of 0.31, with an estimated type B correlation of 0.84, indicating low levels of genotype-by-environment interaction. The trait survival and number of latex vessel rings (RG) showed larger genotype-by-environment interaction and the lowest heritabilites. High to moderate type B correlation was found for most traits, with a value of 0.85 between diameter (or girth) and RYm; therefore, it is possible to achieve interesting rubber yield genetic gains (over 3 years of measurements) from indirect selection based on diameter at age 2.