Number of involved genes and heritability of clover rot (Sclerotinia trifoliorum) resistance in red clover (Trifolium pratense)

European red clover (Trifolium pratense) crops are challenged by clover rot, a devastating disease caused by Sclerotinia trifoliorum or, in some cases by S. sclerotiorum. No completely resistant cultivars are available and resistance breeding is hampered by the lack of knowledge on the number of involved resistance genes and the heritability of clover rot resistance. In this study, we estimated the number of major genes contributing to clover rot resistance by analysing 15 F₁ progeny populations from pair crosses between ramets of resistant and susceptible genotypes. Parent plants were chosen from diverse, diploid populations, including wild material, landraces and cultivars. Young progeny plants were inoculated with ascospores, evaluated phenotypically and the segregation of disease scores was studied. Our results indicated that clover rot resistance may be conferred by three major effect genes, although segregation patterns suggested that there may be numerous minor effect genes involved as well. No proof was found for a maternal inheritance of clover rot resistance. To get insight in the heritability of clover rot resistance, we applied divergent selection by our high-throughput bio-test on an experimental diploid population: the original population (70.5 %), the first generation after selection for susceptibility (79.2 %) and the first generation after selection for resistance (62.3 %) differed significantly in susceptibility (p < 0.001). The second generation after selection for resistance (60.0 %) was not more resistant than the first generation after selection for resistance. In the first generation of selection the heritability (h²) was on average 0.34. In the second generation of selection h² was 0.07. These findings have important implications for resistance breeding.     

Effects of inbreeding on nodal root system morphology and architecture of white clover (<Emphasis Type="Italic">Trifolium repens</Emphasis> L.)

Plants of white clover (Trifolium repens L.) cultivar Crau, a self-fertile Crau genotype, and nine generations of inbred progeny were raised in sand culture in a glasshouse experiment. Digital images of the root systems were made and root morphological characteristics were determined on all the plants. Root architectural parameters were measured on the Crau parent and the S₁, S₄, S₆, and S₉ inbred lines. The clover roots became shorter and thicker with inbreeding but the number of root tips per plant was unchanged. Root architecture (branching pattern) was largely unaffected by inbreeding. It is concluded that inbreeding white clover will lead to shorter, thicker roots, and reduced nutrient uptake efficiency compared with the parent clover. The degree to which these deleterious traits are overcome during the development of F₁ hybrids needs to be determined.     

Resistance to Pseudomonas solanacearum in the potato: II. Aspects of host-pathogen-environment interaction

Summary White clover genotypes were selected for high and low values of leaf size, taproot diamater, number of taproots, proportion of total root and taproot dry weight. Crosses within selected groups of genotypes resulted in 14 seed lines, which were compared with parent genotypes in field tiles. Selections for taproot diameter and proportion taproot were more successful than those for taproot number and proportion root.Narrow sense heritability of characters calculated by regression of progeny on mid-parent values were well correlated with broad sense heritabilities derived from replicated clonal comparisons (r=0.73^*), indicating that the genetic variation is controlled predominantly by additive gene effects.     

Association between growth habit and persistence in red clover

Summary Three double-cut red clover (Trifolium pratense L.) varieties: Prosper I, Florex, and Tristan each with two different seedlots were space-planted in a field. The total number of plants established for each seedlot was about 900. In the seedling year, plants were classified into five growth types according to rosette development and flower production. In the fall of the third year 14% of the rosette-forming, non-flowering Type 1 plants were alive; 8–11% of the intermediate types survived; and only 2% of the nonrosette-forming, profusely flowering Type 5 plants remained growing. The non-flowering class (Types 1 and 2) persisted better than the flowering class (Types 3, 4, and 5). This difference in persistence was similar for all three varieties regardless of seed source. The two flowering classes, in the production years, had the same proportion of the most vigorous plants. The most vigorous plants of the non-flowering class had the same frequency distribution of seed yield as those of the non-flowering class. In conclusion, the flowering response can be used as a selection criterion for persistent plants; and plants not flowering in the seedling year can attain a forage and seed yield as high as flowering plants in the production years. Screening for non-flowering plants can be carried out at the seedling stage in the greenhouse before seedlings are transplanted to the field.Contribution No. 520, from the Charlottetown Research Station, Agriculture Canada, Prince Edward Island, Canada.     

Recurrent selection programs for white clover (Trifolium repens L.) using self-compatible plants. I. Selection of self-compatible plants and inheritance of a self-compatibility factor

Summary When white clover plants were self-pollinated, 75% of a 143-plant population did not set seed. Plants which did set small numbers of seed probably did so as a result of pseudo-self-compatibility. One highly self-compatible plant was selected from the breeding materials. The S₂ progeny of this plant was autogamous and genetically fixed for morphology and isozyme patterns. Genetic analyses of self-compatibility were conducted as far as the F₃ progeny. It was concluded from these data that self-compatibility was inherited in a simple Mendelian way and that plants homozygous for self-compatibility were obtained. There was no correlation between presence or absence of self-compatibility and growth vigor. The possibility of application of self-compatible plants to recurrent selection procedures is outlined and the utilization of self-compatible plants in white clover breeding programs is discussed.     

Effect of various plant factors on the expression of creeping-rootedness in lucerne (Medicago sativa L. complex)

Although agronomically interesting, selection of creeping-rootedness inlucerne, that is, the ability to form adventitious shoots on horizontal,spreading roots, remains difficult because of the complex and still unclear geneticcontrol of this character, coupled with the possible occurrence of external factorsthat affect unpredictably its expression. Two experiments were undertaken onprogenies of creeping-rooted plants, to verify previous findings and inferences,and test novel hypotheses, concerning the effect of various plant factors on theexpression of creeping-rootedness. The ultimate aim was to provide information onthe most appropriate germplasm, procedures, or indirect selection criteria, that wouldimprove the efficiency of breeding for this character. The factors here examinedincluded genotype, age, kind of progeny, vigour, spreading ability, and undergroundmorphology. Some genotypes appeared to be good donors of the character, whereasothers did not. The proportion of creeping-rooted plants increased in allcases with the plant age. Selfing consistently depressed the expression ofthe character compared with clonal propagation or crossing. It is suggestedthat the effect of age and selfing on the penetrance of creeping-rootedness ismediated through their effect on plant vigour. Both aerial and root vigour showedindeed an effect per se on the presence of the character. Beside thevigour, possible indirect selection criteria were identified in the frequenciesof swollen zones and latent stem primordia on horizontal roots. Proportion ofcreeping-rooted plants in progenies after one year of growth appeared as a simple anduseful criterion to predict future trends. Consistent with previous conclusions, theresults provided evidence that a combination of family and individualselection may enhance the character expression.     

Genetic variation in a Panicum coloratum L. population with a limited germplasm base

Summary The U.S. kleingrass (Panicum coloratum L.) germplasm collection is relatively small, and genetic variation for many agronomic traits has not been determined. This study was conducted to explore variability and predict expected progress from selection for several morphological characters related to forage production and establishment in a narrow-based population of kleingrass. Heritabilities and genetic gain were calculated from the analysis of half-sib families in a 2-yr study. Narrow-sense heritabilities were low to moderate and predicted small gains for most characters. Leaf width, however, was consistently heritable in both years, and gains in width of 12.5% per year could be achieved with family selection. Leaf width was also significantly correlated with leaf mass and culm mass. Thus, increases in leaf and culm dry mass accumulation can be achieved by direct selection for leaf width. Two seedling root growth parameters were also heritable, indicating that genetic gain could be achieved for seedling establishment in this population. Heritabilities for number of adventitious roots and length of longest adventitious root at 14 days after emergence were much higher in the second year of the study. This was attributed to the larger number of plants sampled in the second year.     

Heritability and gene effects for root characteristics in peas measured at flowering

Summary Heritability and gene effects for root weight, root volume and root to shoot weight ratio were determined in peas (Pisum sativum L.) at flowering. The populations used were developed from four crosses between lines and cultivars differing in size of the root systems. Broad-sense heritability was between 0.41 and 0.81 for root weight, and between 0.44 and 0.77 for root volume. Additive and dominance effects were important in the genetic control of root weight and volume in all populations, while epistatic effects were important only in two populations. The importance of the genetic parameters in the control of root to shoot weight ratio was unclear. Assuming high correlations between root growth in soil-filled pots in the greenhouse and growth in the field, with the presence of large additive effects and high heritability estimates for root weight and volume, selection for superior pure lines with large root systems should be effective.     

Estimates of heritability for,and relationships between,root and shoot characters of white clover. I. Replicated clonal material

Summary A total of 749 genotypes from a number of white clover (trifolium repens L.) cultivars and populations, many collected from dryland areas, were cloned and grown in field titles. Several morphological characters, including leaf size, number and diameter of large nodal roots (taproots), and proportion of root weight that was classified as taproot were measured. There was large variation between lines and genotypes for all characters measured, and differences between genotypes within lines are also reported. Broad sense heritability estimates were higher (>0.5) for leaflet width, petiole length and stolon diameter, than for all root character heritability estimates which were between 0.2 and 0.4.     

Selection for root hair length in white clover (Trifolium repens L.)

Summary Results are given which show that the root hair length of a white clover population can be increased and decreased by selection. An increase of 50 m in root hair length over the mean of an unselected population of the cultivar Tamar was achieved resulting in an 11% calculated increase in the volume of soil explored by root hairs. The realised heritability for selection in the long root hairs was 0.33, and for the short root hairs 0.44.     

Influence of recurrent selection for flowering on flowering and yields in kura clover

Summary These investigations were conducted to determine the effectiveness of first- and second-season selection programs for increasing forage and seed yield in kura clover (Trifolium ambiguum Bieb.).Rhizo kura clover, the base population, was grown as spaced plants in two programs for six generations of selection for flowering in the seedling and second seasons. Remnant seed for each generation of selection was used to establish spaced plant and broadcast evaluation trials.The first-season selection program was effective in increasing flowering not only in the first season, but for two subsequent seasons. The second-season selection for flowering had little or no effect. However, vigor, forage and seed yields were not increased by either program, and the first-season program progressively reduced vigor and forage yields as generations of selection increased. The cause of this is unknown but may be associated with inbreeding depression or detrimental physiological effects of first-season flowering. Reselection to eliminate possible inbreeding effects is being conducted to further examine the possibility of increasing seedling and aftermath vigor in kura clover.     

土壤水分对冬小麦初生根、次生根生长发育的影响

１９９３－１９９４年冬小麦管栽水分试验结果表明,初生根与次生根生长量,数量及分枝能力基本与土体水量呈正相关,胁迫程度愈深,受抑程度愈大。初生根特别是一,二次根及其分枝数具有很强抗逆逆能力,水量愈大,三,四次根数愈多。次生根尤其是其数量对土水环境极为敏感。当土壤含水量你于田间持水量６０％时,受到严重抑制,胁迫后复水,使初生根三次根数量增加,根系活性延长对次生根的激励作用更大。前期胁迫愈严重,促进效果     

Effects of harvest management on growth dynamics, forage and seed yield in berseem clover

Berseem clover (Trifolium alexandrinum L.) is an annual forage crop widely grown in Mediterranean environments. However, there is little information available on the patterns of accumulation and partitioning of assimilate in berseem and how this varies with harvest management, plant ontogeny or is influenced by genotype. Field experiments were conducted in Foggia (Italy) during three growing seasons (1991–92, 1992–93 and 1993–94), with the aim of evaluating the effects of different harvest managements on growth dynamics, forage yield and seed yield of one population of Egyptian (cv. ‘Giza 10’) as well as Italian (cv. ‘Sacromonte’) origin. Cutting treatments were applied at three different plant ages: sixth internode elongation (A), early flowering (B), and uncut control (C). The growth and development of the shoot and root systems were followed by destructive harvests made at about 5-day intervals during 9 weeks in spring in all treatments. Seasonal growth pattern, determined as dry matter accumulation, forage yield and seed production were greatly influenced by harvest management. The maximum value (20.5 g per plant on average) of dry weight was reached at about 250 days after sowing for treatment C. Defoliation induced a decrease of crown and root dry matter, however, as shown by the allometric relationships, the growth of roots and shoots was closely correlated and in most cases, shoot growth was higher than root growth and stem relative growth rate (RGR) was higher than leaf RGR. Berseem clover is a defoliation-tolerant species because after cutting, regrowth was accompanied by higher values of leaf-stem ratio (LSR), RGR and stem elongation rate (SER) than in control plants. The highest value for total forage yield (1.6 Kg m⁻²) and seed yield (60.5 g m⁻²) were obtained in treatment B and A, respectively. Plants cut at the sixth internode elongation showed a good seed yield and a small decrease (15%) in total forage yield as regards treatment B. Therefore, treatment A appeared the most favourable for obtaining double utilization (forage and seed yield) in a berseem crop. Berseem plants were greatly influenced by harvest treatments applied, but the response did not vary according to the genetic characteristics of the two cultivars examined.     

Endophyte Infection May Affect the Competitive Ability of Tall Fescue Grown with Red Clover

Cool-season grasses infected with Neotyphodium endophytes may be more persistent and competitive than non-infected plants. In a greenhouse experiment, two tall fescue ( Festuca arundinacea Schreb.) cloned genotypes (DN2 and DN11) with different growth characteristics and endophyte status (E−, non-infected; E+, infected) were grown in monocultures and in full competition with red clover ( Trifolium pratense L.) cv. Reddy Red for 20 weeks. When grown in monoculture, endophyte infection reduced the root dry matter (DM) of DN2 (16 %) and DN11 (5 %) when compared to E− plants, while shoot DM was not affected. In full competition with red clover, root and shoot DM of tall fescue were not influenced by endophytes, but cumulative herbage DM yield was less in E+ than in E− plants. Root growth of red clover was significantly depressed (22 %) in competition with E+ plants of DN2 compared to E− plants, but was not affected when plants were grown in competition with DN11. The relative yield total (RYT) did not differ from 1.0 in red clover mixtures with E− plants of DN2 and DN11, and with E+ plants of DN11, indicating competition between the competitors for the same resources. In contrast, RYT was less than 1.0 in the red clover/DN2 E+ mixture, suggesting a negative interaction between the competitors. Endophyte infection increased the competitive ability of DN2 but reduced that of DN11 when compared to E− plants. Because the concentration of pyrrolizidine alkaloids in roots of E+ plants of DN2 was twice that found in roots of DN11 (1083 and 536 μg g⁻¹, respectively), we suggest that pyrrolizidine alkaloids might influence the competitive ability of some endophyte–tall fescue associations through a possible allelopathic effect on companion species.     

Improving seed yield in red clover through marker assisted parentage analysis

Red clover (Trifolium pratense L.) is an important perennial forage crop in Europe. Unfortunately, seed production of modern cultivars is often unsatisfactory and breeding progress for higher seed yield is slow. We evaluated whether marker assisted parentage analysis could identify progeny plants with two high seed-yielding parents in a diploid selection trial. Furthermore, we studied if breeding for high seed yield could be speeded up by selecting progeny plants with two high seed-yielding parents, compared to traditional selection on the mother plant only. In this study, 111 genotypes with excellent vegetative traits from a first cycle selection trial were allowed to pollinate each other in isolation. After seed harvest, ten high seed-yielding plants were identified and their seeds were used to grow ten half-sib progeny populations, with 1,121 plants in total. Two multiplex primer sets targeting 18 SSR loci were designed and used to fingerprint parental and progeny plants. Parentage was reliably determined for 1,083 progeny plants: 135 plants were the result of a cross between two high seed-yielding parents. In a 2-year seed yield trial, the seed yield of these 135 progeny plants was compared to that of a control group of 216 progeny plants from the same ten high seed-yielding mother plants but with no selection on the male contribution, as is current practice. Compared to the control group, progeny plants with two high seed-yielding parents had significantly higher seed yields, 23.0 and 75.9 % higher in 2012 and 2013 respectively, and a significantly higher thousand seed weight, 6.6 and 5.7 % higher in 2012 and 2013 respectively. Seed yield was strongly correlated with flower abundance (r = 0.643) and plant volume (r = 0.593 in 2012 and r = 0.509 in 2013) and negatively correlated with susceptibility to mildew (r = ?0.145). We conclude that breeding programs aiming at increasing seed yield in diploid red clover cultivars would strongly benefit from the integration of marker assisted parentage analysis to identify those progeny plants that are the result of a cross between two high seed-yielding plants. This goal cannot be achieved with traditional family selection where only the seed yield of the mother plant is known.     

Genetic variability and heritability estimates for hardseededness and flowering in balansa clover (Trifolium michelianum Savi) populations

Suitability of annual pasture legume species like balansa clover (Trifolium michelianum Savi) in southern Australian farming systems depends on their hardseededness and time to flowering. Late maturing varieties with increased rate of hardseed breakdown, particularly in the late summer—early autumn period are desirable to ensure reliable regeneration in permanent pastures. Two half-sib family balansa clover populations were used to estimate heritability for the rate of breakdown of hardseededness and time to flowering. High narrow-sense heritability estimates (90.5–96.1%, rate of hardseed breakdown; 85.9–94.5%, time to flowering) were obtained, and were mainly attributed to additive gene effects. There was no relationship between rate of hardseed breakdown and time to flowering in Group1 half-sib family, which indicated that selection for both traits could be undertaken independently. In half-sib family Group 2 the relationship between the two traits suggested the possibility of selecting late maturing cultivars with increased rate of hardseed breakdown. Significant differences in time to flowering and rate of hardseed breakdown both within and between the two half-sib family groups suggested that phenotypic recurrent selection would be effective in improving the traits in these populations.     

Marker‐assisted parentage analysis reveals high individual selfing rates in tetraploid red clover genotypes selected for seed yield

Seed yield is a major breeding target in tetraploid red clover. We investigated if marker‐assisted parentage analysis can identify progeny plants with two high seed‐yielding parents in tetraploid red clover and if this technique is more advantageous than traditional half‐sib selection. Parentage analysis was successfully performed on the progeny from the 10% highest seed‐yielding genotypes from a second‐cycle family selection trial: 16.0% of progeny were identified with a high seed‐yielding father. However, progeny plants with two high seed‐yielding parents did not produce more seeds than traditionally selected progeny (27.3 g vs. 30.7 g/plant, respectively). The 10% highest seed‐yielding genotypes displayed on average 2% self‐fertilization. Four genotypes were self‐fertile with individual selfing rates up to 20%. Our results discourage the use of marker‐assisted parentage analysis to improve seed yield in tetraploid red clover when the material has been preselected for seed yield. Breeders should be aware that intensive selection for seed yield in tetraploid red clover may inadvertently lead to selection for increased self‐fertility, which may increase inbreeding in the long term.     

Morphological characteristics associated with winter survival of five growth types of tetraploid red clover

Summary The aim of this study was to correlate flowering response of tetraploid red clover (Trifolium pratense L.) in the year of seeding with winter survival and other characteristics (vigour, bloom date). In two different years about 2000 plants of two different cultivars (Hungaropoly and Tapiopoly) were established and in the fall were classified into five growth types based on rosette formation and flower production. Non-flowering types were shorter and had few petioles at 6–8 weeks of age, but showed better survival than flowering types. Hungaropoly non-flowering types were more vigorous and bloomed earlier in the second year but Tapiopoly non-flowering types were less vigorous and bloomed later. Although non-flowering in the seeding year was associated with better survival in the second year, there was adequate variation within both types for selection for improved persistence.     

不同生育时期氮素供应水平对杂交水稻根系生长及其活力的影响

在群体水培条件下，以汕优６３为供试材料，于生育前、中、后期设计不同氮素供应水平（１８个处理），研究其对根硪基分化数、不定根数、根重及其根系活力的影响。结果表明（１）生育前期的单株根重随着氮素供应水平的提高而增加，１０、１３、１４、１５叶龄期的单株根重与不定根数均呈极显著正相关，与每条不定根重无显著相关；（２）主茎第１０叶节的根原基分化数与１０叶期的植株含氮率呈正相关（ｒ＝０．９７８＾＊＊），第１０     

甜菜幼苗的分区及其肥大直根的定位研究

本文按照子叶节区理论(Cotyledon node zone theory)研究结果表明, 甜菜幼苗的子叶节区类型为顶枝伸长型(Telome elongation type)。 甜菜肥大直根采用幼苗分区定位法明确其根头部由上胚轴苗区发育而成; 根颈部与根体是由长的顶枝伸长型的子叶节区发育而来; 至于下面短的根尾, 是下胚轴发育的产物; 甜菜真正的根并不肥大