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     

The use of cross prediction methods in a practical potato breeding programme

Summary Most previous studies on cross prediction methods have examined relatively few crosses, particularly in relation to the numbers involved in most breeding programmes. In this paper the feasibility of using cross prediction methods was examined in a practical potato (Solanum tuberosum) breeding scheme by the analyses of progeny from 52 crosses. The variate considered was breeder's preference, a visual assessment made of the harvested tubers to estimate their commercial potential. The results showed that it was possible to identify the superior crosses. Cross prediction based simply on the mean preference scores, averaged over scorers and clones within progenies, estimated on seedlings or first clonal year plants, provided the best estimate of a progeny's performance in the third clonal generation. Predictions based on the expected proportion of clones that would transgress a given target value also provided a good indication of a progeny's potential. The poorest prediction was obtained by using the observed frequency of desirable clones in a progeny sample. The implications for potato breeding are discussed.     

Use of mid-parent values and progeny tests to increase the efficiency of potato breeding for combined processing quality and disease and pest resistance

A potato breeding strategy is presented which avoids the common but ineffective practice of intense early-generation visual selection between seedlings in a glasshouse and spaced plants at a seed site. Once pair crosses have been made, progeny tests are used to discard whole progenies before starting conventional within-progeny selection at the unreplicated small-plot stage. Clones are also visually selected from the best progenies for use as parents in the next cycle of crosses whilst they are multiplied to provide enough tubers for assessment of their yield and quality. Mid-parent values, as well as progeny tests, are then used to select between the resultant crosses. Material from other breeding programmes can be included in the parental assessments and used in the next cycle of crosses if superior. Finally, in seeking new cultivars, the number of clones on which to practise selection is increased by sowing more true seed of the best progenies, but without selection until the small-plot stage. Traits considered are resistance to late blight [ Phytophthora infestans (Mont.) de Bary] and to the white potato cyst nematode [ Globodera pallida (Stone)], fry colour and tuber yield and appearance, as visually assessed by breeders. The theoretical superiority of the strategy for seeking new cultivars lies in being able to practise between-cross selection for a number of economically important traits within 1 or 2 years of making crosses, something that is not possible on individuals as seedlings in the glasshouse or spaced plants at the seed site. This also means that full-sib family selection can be operated on a 3-year cycle, an improvement on current practice of clonal selection on what is often at least a nine-year cycle. New cultivars can be sought with more confidence from the best progenies in each cycle, and modern methods of rapid multiplication used to reduce the number of clonal generations required to find the best clones.     

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.     

Use of tuber progeny tests for genetical studies as part of a potato (Solanum tuberosum subsp. tuberosum) breeding programme

A diallel set of crosses, including selfs and some reciprocal crosses, was made between 15 parents, chosen for their fertility, from those included in a tetraploid potato (Solanum tuberosum subsp. tuberosum) breeding programme at the Scottish Crop Research Institute. The progenies were grown in randomised complete block trials with two replicates at a high-grade seed site from 1994 to 1996 inclusive and at a ware site in 1995 and 1996. The parents were included in the ware trials. Tubers were assessed for visual preference in all trials and for fry colour at both sites in 1996. Emergence and maturity were recorded in the ware trials and the tubers were assessed for yield, dry matter, size, appearance (regularity of shape), scab, uniformity, sprouting in store and keeping quality. There were very few growth cracks and very few internal defects. No reciprocal differences were found. Inbreeding depression was marked for emergence, yield, tuber size and appearance, and visual preference. In contrast, the selfs had a lighter fry colour than the parents and F₁s. Combining-ability analysis (selfs omitted) identified fry colour, emergence, maturity, yield, dry matter and sprouting resistance as traits for which the GCA (general combining ability) variance and narrow-sense heritability were high enough for good progress from full-sib family selection. Correlations between GCAs for pairs of traits were examined, including those from previously published seedling progeny tests. For fry colour, an unfavourable correlation with low yield (r = 0.596) was compensated by a favourable one with high dry matter content (r = 0.652), whereas unfavourable ones between foliage and tuber blight resistance and sprouting susceptibility (r = 0.578 and 0.596) were identified for monitoring. Clones with high GCAs were identified for use as parents in future breeding and the extent to which GCAs could be predicted from the parents, and the offspring means from the midparent means, was determined by regression and correlation analysis. The offspring-midparent regression was highest for fry colour, followed by dry matter, emergence and sprouting. Values were lower for scab due to environmental variation; for maturity, yield and tuber size due to SCA (specific combining ability); and for visual preference due to both factors. The implications for a breeding strategy are discussed. Received: 28 November 1998 / Accepted: 19 December 1998     

Use of seedling progeny tests for genetical studies as part of a potato (Solanum tuberosum subsp. tuberosum) breeding programme

A diallel set of crosses, including selfs and some reciprocal crosses, was made between 15 parents chosen for their male fertility from those included in a tetraploid potato (Solanum tuberosum subsp. tuberosum) breeding programme at the Scottish Crop Research Institute. Seedling progeny tests were used to evaluate the progenies for non-race-specific resistance to late blight (Phytophthora infestans) in both foliage and tubers, quantitative resistance to the white potato cystnematode (PCN) (Globodera pallida) and the commercial worth of their tubers as judged by breeders' visual preference. No reciprocal differences were found. Comparisons of the selfs and crosses revealed inbreeding depression for breeders' preference, which varied among the parents from negligible to severe, whilst there were also statistically significant differences for foliage and tuber blight, but not for PCN. When the selfs were omitted from the combining ability analyses, large differences in general combining ability (GCA) were found for all four traits, and smaller differences in specific combining ability for tuber blight and breeders' preference. The only statistically significant correlation between GCAs for different traits was a favourable one of r = 0.56 between foliage and tuber resistance to late blight. It was concluded that prospects were good for simultaneously improving all four traits by multitrait genotypic recurrent selection.     

Identification of superior parents and crosses in potato breeding programmes

 The effectiveness of parent per se performance and their self values in the selection of superior parents and crosses in potato breeding programmes was studied by evaluating progenies of 72 crosses from 18×4 (female×male) matings, parents and their selfs for ten agronomically important characters for three successive generations. Simple correlation coefficients were computed between parent per se performance versus general combining ability effects, female per se performance versus progeny means of females, female self values versus progeny means of females, mid-parent values versus progeny means of crosses and mid-self values versus progeny means of crosses. The magnitude of the significant correlation coefficients showed that progeny means of crosses could be moderately predicted by the mid-self values for plant vigour and general impression in clonal generations. Similarly, the progeny means of females in clonal generations could be predicted by their per se performance for general impression. Female self values in the second clonal generation were moderately associated with progeny means of females in that generation for general impression and plant vigour. Parent per se performance and mid-parent values were, in general, ineffective in predicting the general combining ability and the progeny means of the crosses, respectively. The poor predictive powers of parent per se performance and mid-parent values are discussed in relation to the relative importance of specific and general combining ability effects. Received: 23 April 1997 / Accepted: 6 August 1997     

The repeatability of progeny means in the early generations of a potato breeding programme

The progenies from eight crosses between potato clones were examined. The clones were grown as seedlings in a glasshouse and subsequently from tubers in the field at two sites in two consecutive years. At harvest, all the clones were visually assessed by four breeders using a 1–9 scale of increasing desirability. Although there was a formally significant interaction between progenies and environments, the rank of the mean of a cross remained relatively consistent over different growing conditions. It is suggested, therefore, that progeny evaluation by breeders' visual preference can be used to identify the crosses with the highest potential of producing commercial cultivars.     

Selection for yield and yield components in the early generations of a potato breeding programme

Summary A number of unselected potato (Solanum tuberosum L.) clones were grown at two locations (a seed site and a ware site) in three consecutive years. The repeatability of total yield and yield components in the first two clonal years was compared with the same characters recorded in the third clonal year. Selection for yield in the first clonal year was only marginally more effective than a random reduction in number of genotypes, while selection in the second clonal year appeared to be somewhat more effective as judged by performance in the third clonal year. The inefficiency of selection in the first clonal year was ascribed, at least in part, to the inaccuracy of yield assessment as well as the carry-over effect of the mother tubers. Correlations of total yield were higher between different years in the same location than between different locations. Selection under growing conditions suitable for production of seed tubers tended to result in selection of early maturing clones which would not necessarily be optimal for ware growing conditions.     

Efficiency of early generation selections for yield and related characters in safflower (Carthamus tinctorius L.)

The efficiency of early generation selection for yield and related characters in safflower (Carthamus tinctorius L.) was studied in the F₂, F₃ and F₄ generations. Twenty-five F₂ progenies derived from various crosses were studied. In the F₂ generation, number of capitula per plant (CNSP), number of seeds per capitulum (SPSP), test weight (SWSP), and seed yield (SYSP) were the criteria used for single plant selection. The analysis of variance showed significant differences for all of the characters in the F₂, F₃, and F₄ generations. The analysis of variance in each of the selection classes showed highly significant genotypic differences. A large number of selections in the CNSP and SYSP classes showed greater yield than the check variety. In each class the mean for that particular character showed a positive shift. The observed F₃ and F₄ means for seed yield per plant was higher in SYSP, indicating the effectiveness of single plant selection for yield. Correlated response showed that selection for number of capitula per plant was effective for improvement of yield.     

Quantitative analysis of the effect of selection history on sugar yield adaptation of sugarcane clones

An objective of the CSR sugarcane breeding programme in Australia was to assess the scope for broadening the genetic base of the commercial sugarcane germ plasm through interspecific hybridization with Saccharum spontaneum clones. The contribution of both selection history and S. spontaneum to sugar yield and its components was investigated in the germ plasm pool assembled. The analysis was conducted on a data-set of 256 clones, consisting of parents and full-sib families generated from 32 biparental crosses, tested in six environments. The minimum number of generations back to S. spontaneum ancestor in the clone's pedigree was used as a germ plasm score. The geographical origin and selection history of each parent and their use in the biparental crosses were used to develop a selection history score for parents and offspring. The variation for seven attributes, cane yield, commercial cane sugar %, sugar yield, stalk number per stool, stalk weight, fibre % and ash % juice was partitioned according to the germ plasm and selection history scores. Significant (P<0.05) clone variation and clone x environment interaction for all attributes was present. The germ plasm scores accounted for a significant (P<0.05) component of the clone variation for all of the attributes except cane yield. There was an increase in sugar yield with an increase in the minimum number of generations back to a S. spontaneum clone. The selection history groups accounted for a high proportion of the variation among parental clones for all of the attributes except cane yield. This suggested that parents were the outcome of strong selection pressure for the commercial cane attributes. However, the selection history groups for the offspring produced by random mating of parents did not account for a high proportion of the variation for the attributes. Using the mixture method of classification we partitioned the 256 clones into five groups for patterns of performance for the seven attributes across the six environments. The five groups emphasized major differences in the patterns of performance for the seven attributes across environments. The distribution of germ plasm and selection history scores in each of the five groups indicated that their patterns of performance were associated with selection history and minimum generations to S. spontaneum. Therefore, both the analysis on selection history and germ plasm scores (extrinsic classification) and the analysis on the mixture method of classification (intrinsic classification) emphasized the influence of selection history on the sugar yield of sugarcane.     

Breeding of nodulated red clover (Trifolium pratense) for high yield

The symbiotic effectiveness of four cultivars of red clover were compared and breeding programmes undertaken to increase nitrogen fixation and yield of agar- and pot-grown plants. Programme I used the moderately effective Rhizobium trifolii strain 0403 and Programme II the highly effective strain 5. Aggregate scores of plant size (leaf area) were chosen as the criterion of selection. All cultivars, inoculated with strain 0403, differed in time of initial nodulation and in dry matter yield and three differed in leaf area. None differed in nodule number or N-content. Dry matter and leaf area were highly correlated. Diallel crosses among highly effective selections in Programme 1 gave progeny yields that exceeded those of crosses between modal selections by averages of 6% in the first generation, 5% in the second generation and 23% in the third generation. Yields of the high cross category exceeded those of the original cultivar by an average of 9% in the second generation and by 25–101% in the third generation. Effects on leaf area were similar. Highly effective progeny tended to nodulate earlier and have more nodules than the remainder but differences were very small. Crosses between cultivars were generally more effective than those within cultivars, indicating heterotic effects. Similar results for yield were obtained in Programme II. The average increase in yield of crosses of third-generation material between highly effective selections compared with cv. S123 was 63%. The modal crosses were intermediate. In each Programme and generation there were large differences between parents within each cross category.     

The efficiency of visual selection in early generations of a potato breeding programme

The efficiency of visual selection in the early generations of a potato breeding programme is examined. Tubers from 571 potato genotypes were scored by four breeders after being grown, from true seed, in a glasshouse and in the field for three consecutive years. The four breeders were in good agreement as to which clones would be selected in any environment. The association between breeders was greatest when the clones were grown in plots in the field. All correlations of breeders' preferences between different environments were significantly greater than zero, but only accounted for a small percentage of the total variation. Selection in both the glasshouse and first clonal year produced a desirable response. However, such selection carried a high cost in terms of losing clones with commercial potential. Comparison of a random sample of clones with ones from the same crosses which had been selected, indicated that selection in the glasshouse and first clonal year was at best random with some suggestion, however, of a negative effect.     

Spatial variation in local pollen flow and mating success in a Picea abies clone archive and their implications for a novel ��breeding without breeding�� strategy

Currently, Norway spruce (Picea abies) breeding in Sweden is based on crosses between the best clones followed by clonal testing of the progenies to select for the long-term breeding population. An alternative breeding strategy called “Breeding without Breeding” (BwB) is proposed, which, in principle, relies on the DNA marker-based pedigree reconstruction from wind-pollinated progenies instead of controlled crosses. To test whether the pedigree structure could be established from progenies of clonal trials, we investigated the spatial pattern of local pollen flow and paternity assignment in a clone archive of Norway spruce. The results showed that 42% of the progeny can be assigned to fathers within 30-m distance with high confidence. Effective pollen dispersal decreased rapidly with distance and followed exponential distribution on local scale. The extent of close-neighbor (within 6 m) mating ranged from 0% to 48% among grafts with an average of 13%. Distance explained 25% deviance in mating success, and other factors such as phenology and spatial configuration of the clones should have contributed the rest. The success of parentage assignment in clone archive opens up the possibility to apply BwB in clonal trials of species that are easy to propagate vegetatively. This procedure could substantially shorten the breeding cycle and still give similar gain per year as the conventional breeding.     

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     

Multitrait selection system using populations with a small number of interploid (4x-2x) hybrid seedlings in potato: Degree of high-parent heterosis for yield and frequency of clones combining quantitative agronomic traits

One of the most attractive features of the 4x-2x breeding scheme via unilateral sexual polyploidization (USP) is the possibility of obtaining heterotic clones with a combination of desirable traits based upon selection in a small number of hybrid seedlings. A set of experiments was carried out to verify this characteristic of the USP scheme using 42 families (with 20 plants each) derived from 2x Phureja-haploid Tuberosum or haploid Tuberosum-Solanum chacoense hybrids as male parents. These clones were 2n-pol-len-producers by either first-division restitution with crossing over (FDR-CO) or without crossing over (FDR-NCO). The 4x parents were eight cultivars of USA and European origin. A total of 168 out 840 clones was initially selected (four clones/family) based on general tuber appearance (GTA). An additional round of selection (not taking into account the parentage of the clone) for total tuber yield (TTY) reduced the sample to 96 clones. These selected clones and the original 4x parents were evaluated in two locations at Wisconsin (Hancock – E?1, and Rhinelander – E?2). The average high-parent heterosis values for TTY ranged from 27.8% (E?2) to 48% (E?1). The population of experimental clones also had a TTY range wider than that of the 4x parents. The best clone gave a yield of 101.6% and 63.2% over the best 4x parent at E?1 and E?2, respectively. For TTY, a significant genotype×environment (G×E) interaction for the experimental clones was found when a combined analysis of variance for both locations was carried out. However, the G×E interaction was not significant when only the 4x-parent group was considered. Clones derived from 4x European cultivars had higher GTA scores than clones derived from 4x USA cultivars. With two culling levels being set on the TTY≥4x parent group mean and the GTA≥three, about 56% (E?1) and 48% (E?2) of the clones would be retained for further evaluation. These percentages of selected clones are much higher than those reported using conventional 4x-4x crosses. Our results indicate that a USP strategy with 4x-2x (FDR) crosses would be more effective than intra-Tuberosum crosses in generating heterotic clones with a combination of desirable quantitative traits using populations with a small number of hybrid seedlings.     

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.     

Direct estimation of the effects of meiotic recombination on potato traits via analysis of 4x-2x progenies from synaptic mutants with 2n-pollen formation by FDR without crossing-over

Diploid potato clones with 2n-pollen formation by first-division restitution without crossing-over (FDR-NCO) are ideal testers to estimate the breeding value of elite 4x cultivars by virtue of transmitting their genotypes practically intact to their progenies. This characteristic facilitates genetic analysis, since meiotic recombination would take place only in the 4x parent and not in the diploid parent. We evaluated (under short-day conditions) families from complete factorial crosses between four 4x cultivars and five 2x(FDR-NCO) clones. Families were compared with two standard 4x cultivars (’Bintje’ and ’Delta’) for total tuber yield (TTY), commercial yield (CY), haulm maturity (HM), plant vigor (PV), plant-top uniformity (PU), eye depth (ED), number of tubers per hill (NTH), and the CY/TTY index (CTI). For TTY, the contrasts family group (310 g/ hill) vs ’Delta’ (430 g/hill) and the family group vs ’Bintje’ (210 g/hill) were significant. Only 25% of the families were different from ’Delta’ and 20% of them outyielded ’Bintje’. For CY, differences were observed between families (240 g/hill) vs ’Delta’ (340 g/hill) and families vs ’Bintje’ (150 g/hill). The two best families had 53% CY over ’Bintje’. Surprisingly, only one family had a higher NTH than ’Bintje’. No differences were observed for HM. Seventy five and 30% of the families had an ED similar to ’Delta’ (ED = 2) and ’Bintje’ (ED = 1), respectively. A multivariate analysis indicated that 63% of the data variability could be explained by two factors. TTY, CY, and PV had high loading on the first factor, whereas ED, PU and HM had high loading on the second factor; CTI and NTH had equal sizes on both factors. High TTY and PV were associated with high NTH and CTI. Deep eye, PU, and late maturity were associated with high NTH and reduced CTI. The distributions of factor scores of the entries indicated that some 2x parents had strong influences (irrespective of the direction of their effects) on the crosses. Six crosses due to two 2x males were in the ’Bintje’ quarter with negative scores for both factors (implying low TTY, poor vigor, and low NTH). Also three crosses due to another 2x clone were distributed in the quarter of positive factor 1 and negative factor 2. These crosses plus another one were in the same quarter of ’Delta’ (implying high yields, low ED, low PU, and early maturity). The FDR-NCO clones provide a homogeneous sample of heterozygous 2n-gametes allowing the unique opportunity to estimate the relative contribution of the random meiotic products (from the 4x parents) and the ’somatic’ 2x genome for the phenotypic expression of quantitative traits. The interesting result was that measurable effects (favorable or not) on the data variability were mainly determined by the genomic contribution of the haploid-species hybrids. Three out of five 2x-male parents showed rather strong effects on progenies. No such effects were observed on the four 4x-female parents. Received: 3 September 1999 / Accepted: 24 November 1999     

Experimental life-history evolution: selection on the allocation to sexual reproduction and its plasticity in a clonal plant

Abstract.— Allocation to sexual reproduction is an important life-history trait in clonal plants. Different selection pressures between competitive and competition-free environments are likely to result in the evolution of specialized genotypes and to maintain genetic variation in reproductive allocation. Moreover, selection may also result in the evolution of plastic allocation strategies. The necessary prerequisite for evolution, heritable genetic variation, can best be studied with selection experiments. Starting from a base population of 102 replicated genotypes of the clonal herb Ranunculus reptans , we imposed selection on the proportion of flowering rosettes in the absence of competition (base population: mean = 0.391, broad-sense heritability = 0.307). We also selected on the plasticity in this trait in response to competition with a naturally coexisting grass in a parallel experiment (base population: 14% lower mean in the presence of competition, broad-sense heritability = 0.072). After two generations of bidirectional selection, the proportion of flowering rosettes was 26% higher in the high line than in the low line (realized heritability ± SE = 0.205 ± 0.017). Moreover, genotypes of the high line had 11% fewer carpels per flower, a 22% lower proportion of rooted rosettes, and a 39% smaller average distance between rosettes within a clone. In the second experiment, we found no significant responses to selection for high and low plasticity in the proportion of flowering rosettes (realized heritability ± SE =–0.002 ± 0.013). Our study indicates a high heritability and potential for further evolution of the proportion of flowering rosettes in R. reptans , but not for its plasticity, which may have been fixed by past evolution at its current level. Moreover, our results demonstrate strong genetic correlations between allocation to sexual reproduction and other clonal life-history characteristics.     

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