Selection of Diploid Nematode-Resistant Sugar Beet from Monosomic Addition Lines

Monosomic sugar beet/wild beet addition lines (2n = 19) with full resistance against the beet cyst nematode have been characterized in different ways. Within the B. procumbens and B. webbiana addition lines three groups could be classified according to their isozymes pattern, growth habit, transmission rate, and resistance level. It is assumed that B. procumbens and B. webbiana each possess three different chromosomes which carry genes for nematode resistance. In the offspring of the addition lines diploid translocation types appear at very low frequencies, Isozyme pattern or growth type of the resistant plants were used for selecting diploid types in the offspring of monosomic addition lines. Effective selection could be made in progenies of susceptible sugar beets pollinated by addition lines because the pollen transmission of the alien chromosome is very low. Using these methods 7 nematode-resistant sugar beet lines could be selected. The transmission rates of the resistance gene ranged from 70.6% to 100%. Threw heterozygous progenies showed a 1:1 segregation indicating monogenic dominant inheritance of resistance. The level of resistance was as high as in the addition lines.     

The production of alien monosomic additions in Beta vulgaris as a source for the introgression of resistance to beet root nematode (Heterodera schachtii) from Beta species of the section Patellares

Summary Experiments were carried out for adding the chromosome carrying resistance to beet root nematode (Heterodera schachtii) from the wild Beta species of the section Patellares (B. procumbens, B. webbiana and B. patellaris) to the genome of B. vulgaris. Preliminary experiments indicated that crosses between the wild species and B. vulgaris cultivars of the mangold type yielded on average more viable F₁ hybrids than crosses with sugar and fodderbeet. However, crossability varied strongly between individual parental combinations. It was concluded that most types of B. vulgaris can be hybridized with the wild species of the section Patellares if a sufficient number of pair-crosses is made. Crosses between diploid cultivars or species of the section Vulgares and diploid wild species of the section Patellares yielded many hybrids which, however, were highly sterile. From crosses between tetraploid B. vulgaris and the wild species a great number of viable allotriploid and allotetraploid hybrids was obtained. In the backcross progenies of allotriploid hybrids 26% alien monosomic additions occurred, of which 4.1% carried the resistance bearing chromosome of B. procumbens or B. patellaris. The programme will be continued by sereening progenies of the resistant monosomic addition plants for the occurrence of resistant disomic introgression products.     

Resistance to Heterodera schachtii in patellares section of the genus Beta

Summary Fifty-two accessions of the section Patellares wild beet (including 26 Beta patellaris Moq., 13 B. procumbens Chr. Sm and 13 B. webbiana Moq.) and 14 progeny families were selected and tested against sugarbeet cyst nematode, Heterodera schachtii Schm. All Patellares species tested were highly resistant, but not immune, to the development of H. schachtii, after infection. This is the first report that mature female nematodes developed in the roots of B. webbiana plants. The occasional development of nematode cysts in roots of juvenile wild beets was, however, not a heritable genetic factor.Mention of a trademark, vendor, or proprietary product does not constitute a guarantee or warranty of the product by the USDA and does not imply its approval to the exclusion of other products or vendors that may also be suitable.     

Partial resistance of sugarbeet to beet cyst eelworm (Heterodera schachtii Schm.)

Summary Within Beta vulgaris and B. maritima origins some partial resistance to beet cyst eelworm was found which could be raised to a very limited extent by selection. However after the second backcross to commercial sugar beet varieties and successive selection of the inbreds this resistance was lost. It was demonstrated that in the rootsystem of resistant plants as much nematodes penetrate and develop as in susceptible ones but the ratio between males and females is different. It was therefore quite probable that this resistance is polyfactorial and merely recessive.     

Expression in vitro of resistance to Heterodera schachtii in hairy roots of an alien monotelosomic addition plant of Beta vulgaris,transformed by Agrobacterium rhizogenes

Summary Hairy roots, induced by Agrobacterium rhizogenes, were obtained of nematode susceptible beet plants (Beta vulgaris) and of the nematode resistant alien monotelosomic addition AN5, carrying a telosome from B. patellaris. The additional telosome was found to be stably present in vitro in the roots of AN5. The hairy root cultures were inoculated with larvae of the beet cyst nematode Heterodera schachtii. On the root culture of AN5 significantly less cysts developed than on the other root cultures. These results indicate that the resistance to the beet cyst nematode is expressed in the roots after transformation and can be monitored under in vitro conditions.     

Nematode Resistance Derived from Wild Beet and its Meiotic Stability in Sugar Beet

Three different karyotypes of sugar beet with resistance against the beet cyst nematode (Heterodera schachtii) have been investigated. These comprised monosomic addition lines (2n = 19) with one complete chromosome from B. patellarris or B. procumbens, one line with a chromosomal fragment added to the normal sugar beet chromosome complement (2n =18 + fragment) and one diploid line (2n = 18). The fragment originated from a B. procumbens chromosome since during meiosis it formed a univalem. It carries the gene for nematode resistance. Meiotic disturbances like univalems. laggards, anaphase I bridges, fragments and micronuclei were observed in all resistant genotypes. These may result in an exclusion of the chromosome fragment carrying the resistance from the rest of the genome. In the diploid resistant line, a chromosome with a translocation could be distinguished from the other B. vulgaris chromosomes. Meiotic irregularities also appeared in diploid resistant types and are one main reason for low transmission of the resistance. Tin-relationship between meiotic stability and the transmission rate of the resistance gene is discussed.     

Sources of resistance to diseases of sugar beet in related <Emphasis Type="Italic">Beta</Emphasis> germplasm: I. Foliar diseases

Resistance to four foliar diseases of sugar beet (Beta vulgaris ssp. vulgaris), virus yellows caused by Beet mild yellowing virus (BMYV) and Beet yellows virus (BYV), powdery mildew (Erysiphe betae) and Cercospora leaf spot (Cercospora beticola), was assessed in up to 600 accessions of closely related wild and cultivated Beta species. Most accessions were from the Section Beta, a taxon containing types most closely related to, and sexually compatible with, sugar beet and therefore most valuable for use in crop improvement. Between 1–12% of accessions were highly resistant (resistance scores of 2 on an international standardised resistance scale of 1–9) to these diseases. These levels, however, underestimate the potential number of resistant sources available from this section as some accessions with intermediate mean resistance scores contained a significant proportion of highly resistant plants within segregating populations. Variation in resistance to all diseases except BYV was observed within the Section Beta. Much higher levels of resistance were observed, and more frequently, in more distantly related sections of the genus Beta. Accessions of the Section Corollinae were highly resistant to both viruses (>62% of accessions tested), but less so to Cercospora (15%) and they were very susceptible to powdery mildew. Section Procumbentes accessions were highly resistant to BMYV and Cercospora (100%) but less so to powdery mildew (50%) and BYV (20%). However, sexual incompatibility between these sections and sugar beet make utilisation of these sources impractical using conventional breeding methods.     

Major genes for resistance to beet necrotic yellow vein virus (BNYVV) in Beta vulgaris

Summary Inheritance of resistance to beet necrotic yellow vein virus (BNYVV) was studied in segregating F2 and backcross families obtained from crosses between resistant plants of the sugar beet selection Holly-1-4 or the wild beet accession Beta vulgaris subsp. maritima WB42 and susceptible parents. Greenhouse tests were carried out, in which seedlings were grown in a mixture of sand and infested soil. Virus concentrations of BNYVV in the rootlets were estimated by ELISA. To discriminate resistant and susceptible plants, mixtures of normal distributions were fitted to log₁₀ virus concentrations, estimated for segregating F1, F2 and BC populations of both accessions. The hypothesis that Holly-1-4 contained one single dominant major gene was accepted. For WB42, results fitted with the hypotheses that resistance was based on either one (or more) dominant major gene(s) showing distorted segregation, or two complementary dominant genes, which are both required for resistance. Resistance from WB42 appeared to be more effective against BNYVV than resistance from Holly-1-4.This research was carried out as part of a PhD study at the Graduate School Experimental Plant Sciences (EPS), Department of Virology, Wageningen, The Netherlands     

Beet mild yellowing virus resistance derived from wild and cultivated Beta germplasm

Sugar beet (Beta vulgaris ssp. vulgaris L.) has a relatively narrow genetic base and several programmes have sought to evaluate the potential for introducing novel traits from wild and cultivated Beta germplasm into the crop. In particular, resistance to important sugar beet diseases has been identified within individual Beta accessions. We report here the successful transfer of resistance to Beet mild yellowing virus (BMYV) from garden beet, fodder beet and leaf beet accessions to progeny populations in initial crosses with sugar beet. Twelve plant populations derived from different Beta accessions were inoculated with viruliferous aphids carrying BMYV and the virus content of individual plants subsequently quantified by an ELISA test. Seven populations were significantly more resistant than a control sugar beet cultivar (P ≤ 0.05). BMYV resistance was successfully inherited in BC₁ and BC₂ generations, suggesting that resistance could potentially be introgressed from these sources into elite sugar beet lines.     

Enhanced resistance to a lepidopteran pest in transgenic sugar beet plants expressing synthetic <Emphasis Type="Italic">cry1Ab</Emphasis> gene

Two diploid sugar beet genotypes of agronomical importance were transformed using Agrobactrium tumefaciens harboring pBI35Scry containing a synthetic cry1Ab gene. Leaf blade with attached shoot bases, a highly regenerative tissue, were used as explant substratum for transformation. PCR screening with cry1Ab-specific primers showed the presence of transgene in more than 50% of the regenerated kanamycin-resistant plants after treatment with the antibiotic. A transformation rate of 8.8–12.2% (depending on genotype) was achieved as revealed by genomic DNA dot blotting. The intact integration of transgene cassette into the genome was furthermore confirmed by Southern blot analysis. The expression of the cry1Ab gene encoding a truncated endotoxin (67 kDa) at about 0.1% of total soluble protein was achieved in the leaves of transgenic plants as shown by Western blot analysis. Bioassays under in vitro conditions with Spodoptera littoralis, one of the most important pests in sugar beet fields, demonstrated enhanced resistance against this pest. The inheritance of the inserted transgene was confirmed in F₁ plants obtained through crossing of T₀ plants with a cytoplasmic male sterile line. Transgenic plants are currently grown in a greenhouse and will be subjected to further bioassay analyses against other lepidopteran pests of sugar beet.     

Breeding for resistance to rhizomania in sugar beet: A review

Currently rhizomania is the most important disease in sugar beet worldwide, and attack can lead to serious yield losses. The disease is caused by beet necrotic yellow vein virus (BNYVV) that is transmitted by the soil-borne fungus Polymyxa betae. Breeding sugar beet cultivars with resistance to rhizomania is regarded as the most appropriate way to enable continued production of this crop in BNYVV-infested fields and also to slow the spread of the disease. Breeding for resistance started with selection by scoring disease symptoms in field experiments. The development of non-destructive greenhouse tests, with determination of the virus concentration in rootlets using ELISA, has greatly improved the efficiency of selection. In this paper the impact of scientific research on the progress in breeding cultivars with resistance to rhizomania is reviewed. This includes the distribution, composition, and pathogenicity of the virus, the sources of resistance to virus and vector, the genetics of virus resistance, progress with breeding methods, and the use of molecular markers and pathogen-derived resistance. The yields and quality characteristics of recently introduced resistant cultivars now equal those of the commercial susceptible cultivars. This revised version was published online in July 2006 with corrections to the Cover Date.     

Greenhouse and field techniques for testing sugar beet for resistance to Rhizoctonia root and crown rot

Rhizoctonia‐resistant sugar beet varieties are the key to an integrated control strategy for Rhizoctonia root rot. Because of the unpredictable occurrence of Rhizoctonia solani in the field testing of sugar beet for resistance to Rhizoctonia root rot is difficult. The aim of the study was to develop advanced greenhouse and field techniques which allow a reliable assessment of sugar beet for resistance to R. solani. A highly infectious liquid inoculum was used for the first time in this study. It can be produced in large quantities of a standardized quality, sterile, and exactly quantified according to its carbon content. In a greenhouse trial, sugar beet grown in the same way as field grown beet was inoculated by applying a suspension of Rhizoctonia mycelium (equivalent to 10 mg carbon per plant) to the beet crown. After 3 weeks, inoculation had led to uniform and severe root rot. Disease symptoms were similar to those found under natural conditions in the field. No ‘escapes’, i.e. susceptible plants apparently expressing resistance were observed in the test. A new nine‐class disease scale was established and a Rhizoctonia index (RI) was calculated. Reliability of disease assessment was demonstrated on progeny of plants, selected from segregating populations, showing Rhizoctonia resistance more closely related to the resistant parent lines than to the susceptible ones. Sugar beet varieties could be assessed in the greenhouse within only 11 weeks. All varieties were affected by the pathogen but partially resistant varieties could be clearly recognized by a significantly lower Rhizoctonia index. Significant differences in susceptibility were also found within the group of new resistant genotypes. Infection studies performed in the field showed the superiority of the new liquid inoculum compared with a solid form and revealed the influence of inoculation date and inoculum level on the development of Rhizoctonia root rot. In field tests performed at different sites under different environmental conditions, susceptible and partially resistant sugar beet varieties could be reproducibly rated according to their susceptibility to R. solani. On average, susceptible varieties showed a Rhizoctonia index of 8 while resistant genotypes ranged from 5 to 6. The newly developed techniques allow fast and reliable evaluation of sugar beet for resistance to R. solani.     

Assessment of the degree of genetic variation in beet based on RFLP analysis and the taxonomy of Beta

Summary Clones derived from Beta vulgaris and Beta maritima were assayed for their ability to detect restriction fragment length polymorphisms (RFLPs) in different beet accessions. The clones able to detect polymorphism were used as genetic markers to assess the degree of genetic variation existing between and within different species of the genus Beta. The data support the current taxonomy of the Beta vulgaris section, while the great genetic similarity found between Beta webbiana and Beta procumbens indicates that they could belong to the same species.Enough variation was found between Beta vulgaris cultivars, allowing the isolation of a sufficient number of genetic markers for the construction of detailed genetic maps.     

Sources of resistance to diseases of sugar beet in related Beta germplasm: II. Soil-borne diseases

Between 580 and 700 accessions of related cultivated and wild species of the genus Beta were assessed for resistance to four soil-borne diseases of sugar beet: two seedling damping-off diseases caused by the fungi Aphanomyces cochlioides and Pythium ultimum and two diseases of more mature plants, Rhizoctonia root and crown rot, caused by the fungus R. solani, and Rhizomania, caused by Beet necrotic yellow vein virus (BNYVV), a furovirus transmitted by the plasmodiophorid Polymyxa betae. Analysis of resistance data (assessed on an international standardised 1–9 scale of Resistance Scores) indicated that the highest levels of resistance ({RS} 2) to A. cochlioides and P. ultimum were to be found amongst accessions of the more distantly related sections Corollinae (93% of accessions tested) and Procumbentes (10%), respectively; although useful levels could also be found in the more closely related, and sexually compatible, section Beta (1–6%). Resistance to Rhizoctonia was also found in section Beta (5–7%), depending on whether field or glasshouse tests were used, but there was little evidence of generally high levels of resistance to Rhizomania among accessions of this section. None of the accessions of sections Corollinae and Procumbentes exhibited any notable resistance to Rhizoctonia. However, all sections Procumbentes and some sections Corollinae (4%) accessions were highly resistant to Rhizomania. Individuals with high levels of resistance to Rhizomania were identified from within some section Beta and Corollinae accessions, in which there was evidence of segregation.     

Nematode Resistance and Meiotic Abnormality in Diploid Sugar Beet Selected from Interspecific Beta vulgaris×B. procumbens Hybrids*

Two nematode-resistant trisomic lines which were derived from interspecific Beta, vulgaris × B. procumbens hybrids were intercrossed or backcrossed with susceptible diploid sugar beer and their progenies were screened for nematode resistance. The transmission rate of resistance varied from 1.5 % to 47.6 % with an average of 20.4 % in the progenies of individual insomics derived from the two trisomic lines. Eleven resistant diploads were selected with a frequency of 0.2 %. These resistant diploids were classified into two groups, i.e., one group showed relatively high transmission rates of resistance with an average of 25.4 % and the other extremely low with an average of 1.2 % in their backcrossed and s el fed progenies., Meiotic chromosome behavior in a resistant diploid group with high transmission rates was considerably normal as compared to that in a resistant diploid group with low transmission rates. Chromatid bridges and acertric fragments were detected in 93 % of resistant diploids and in 46 % of susceptible diploids. Two different sized fragments occurred in resistant diploids, while only a smaller fragment was present in susceptible diploids. A frequency of sporocytes with bridges-fragments was 17.4% at anaphase I and 13.9 % at anaphase II in resistant diploids, while in susceptible diploids a frequency was 2.9 % and 5.3 % at the respective stages. These results suggest that at least two paracentric inversions are present in resistant diploids, one of which is linked to nernatode resistance and may be responsible for the low transmission rate of resistance.     

Origin of the ‘Weisse Schlesische Rübe’ (white Silesian beet) and resynthesis of sugar beet

Summary Sugar beet-besides fodder beet, red beet, and chard-belongs to Beta vulgaris L. After it had been confirmed that the sugar of Beta beet is chemically identical with cane sugar, ACHARD started experiments on the production of sugar from fodder beet. He noticed that conical white beets seemed to have the highest sugar content. This first sugar beet, the Weiße Schlesische Rübe, is considered the ancestor of all sugar beets of today. It has been, and continues to be supposed that it had originated from crossings between typical fodder beet and chard. Hints in the literature about possibilities to resynthesize sugar beet by crossing fodder beet with chard were confirmed in the author's own trials; the F₂ from the crossing fodder beet Rote Walze x chard Lukullus segregated forms and colour variants largely corresponding to sugar beet. Such new sugar beets are not only important from a theoretical point of view; breeders are interested in new types, too. The synthesis of sugar beet is interpreted from a genetic point of view.     

Ecological aspects of transgenic sugar beet: transfer and expression of herbicide resistance in hybrids with wild beets

Summary An increasing number of genetically engineered cultivars of several crops is being experimentally released into the environment. In future, crops with new transgenic traits will probably play an important role in agricultural practice. The long-term effect of transgenes on community ecology will depend on the distribution and establishment of transgenic plants in the wild, on the sexual transfer of their new genes to the environment and on the potential ecological impact of the transgenic trait. The starting point was the use of transgenic sugar beet lines, Beta vulgaris subspec. vulgaris var. altissima DÖLL (Helm 1957), with transgenes coding for rhizomania and herbicide (BASTA^®) resistance. The first two questions to answer were: Can the transgenes be transferred via pollen to wild beets, Beta vulgaris subspec. maritima (L.) ARCANG. or cultivated relatives such as red beet or spinach beet and are they expressed in the hybrids? Can transgenes be monitored in young Beta vulgaris-hybrids? The experimental transfer of transgenes was conducted in 1993 at a field location in northern Germany. The beets were hand-pollinated with transgenic pollen. In a non destructive biotest, the hybrid seedlings were tested for herbicide resistance. Transgenic plants showed no noxious phenotypic effects whereas control plants developed leaf necroses. All herbicide resistant hybrids within the biotest were assumed to be transgenic.     

Electrophoretic Investigations on Nematode Resistant Sugar Beets

Hybridizations were made between beta vulgaris and three wild species of the patellares section being resistant to the best cyst nematode {heterodera schachtii). Monosomic addition lines (2 n = 19) with full nematode resistance were investigated together with wild beets by means of electrophoretical techniques. One alkaline esterase band and a complex of several acidic esterase bands were localized on the resistance-carrying B. procumbens chromosome. The alkaline esterase marker also appeared in B. patellaris addition lines. An aconitase double band was visible in two of four B. webbiana addition lines. One resistant monotelosomic addition line with a small B. procumbens fragment had lost the esterase gene. Evidence is given that more than one chromosome is carrying genes for nematode resistance. The use of electrophoretic screening together with a nematode testing program is discussed.     

Peroxidase activity as a possible test for distinguishing s-plasm from n-plasm in sugar beet (Beta vulgaris L.)

Summary The O-type lines (N-plasm) of sugar beet (Beta vulgaris L.) proved to have a consistently lower peroxidase activity (49.2–62.7%) than their male sterile (MS) equivalents (S-plasm).Similar differences in peroxidase activity (45.4–56.3%) were found when O-type lines were compared with unrelated MS lines. They were also observed in different parts of the plant (cotyledons, hypocotyl + radicle, true leaves) and were reproducible.The fertile lines could be grouped by their level of peroxidase activity in categories of either S-plasm or N-plasm.The peroxidase activity test could be used for selection of N-plasm lines from fertile sugar beet populations and for selection of MS and O-type lines of higher stability.