Inheritance of resistance to head bugs and its interaction with grain molds in Sorghum bicolor

Sorghum head bug, Calocoris angustatus Lethiery is one of the most important pests of grain sorghum in India. Head bug damage increases the severity of grain molds, which renders the grain unfit for human consumption. Therefore, we studied the gene action for resistance to head bugs and grain molds in a diverse array of male-sterile lines and testers in a line × tester mating design under natural infestation. Mean squares for parents, parents vs crosses, lines, testers, and lines × testers were significant for head bug damage and grain mold severity. General combining ability (GCA) effects were significant and negative for ICSA 88019 for head bug damage, and ICSA 88019 and ICSA 88020 for grain molds (except for ICSA 88020 in 1993). General combining ability effects were positive for ICSA 42 and 296 A. GCA effects of lines and testers for head bug damage and grain mold severity were in the same direction (+ve or −ve). Head bug damage in the grain was significantly correlated with grain mold severity. Testers IS 8891, IS 15107, and TAM 2566 (with colored grain and less susceptibility to molds) produced mold-resistant hybrids in combination with all the male-sterile lines, while the reverse was true in the case of Swarna and ICSV 112. Resistance to head bugs showed dominance to partial dominance type of gene action, while in the case of grain molds, it showed dominance to over dominance. Resistance to these pests is governed by both additive and nonadditive types of gene action. The implications of these results are discussed in relation to need for crop improvement in sorghum. This revised version was published online in July 2006 with corrections to the Cover Date.     

Mechanisms and diversity of resistance to sorghum midge, Stenodiplosis sorghicola in Sorghum bicolor

Sorghum midge, Stenodiplosis sorghicola (Coquillett) is the most important pest of grain sorghum worldwide, and plant resistance is an important component for the control of this pest. To identify sorghum genotypes with diverse mechanisms of resistance to sorghum midge, we studied oviposition, larval survival, and midge damage in 27 sorghum midge-resistant genotypes, and a susceptible check under greenhouse conditions. Observations were also recorded on floral characteristics and compensation in grain mass. Of the 28 sorghum genotypes tested, 19 showed high levels of antixenosis to oviposition as a component of resistance, and had <20% spikelets with eggs when infested with 10 or 25 sorghum midge females per panicle under no-choice conditions in the headcage. Genotypes IS 8887, IS 10712, IS 21873, IS 21881, ICSV745, and QL 39 showed antibiosis as one of the components of resistance. Lines IS 7005, IS 10712, IS 18563, IS 21873, IS 21881, PM 15936-2,ICSV 197, and ICSV 745 showed <20% spikelets with eggs, larvae,or, midge damaged chaffy spikelets across infestation levels, compared with >80% midge damaged spikelets in QL 12 - the susceptible check. Genotypes showing resistance to sorghum midge have smaller glumes than the susceptible check, QL 12. However, IS 7005, IS 18653, and ICSV745 have relatively large sized glumes, but suffered <20% midge damage suggesting that factors other than glume size also contribute to midge resistance in sorghum. Fourteen genotypes showed >20% compensation in grain mass when the panicles were reduced to 250 spikelets and infested with 10 or 25 midges per panicle. There is considerable diversity in sorghum genotypes showing resistance to sorghum midge. Genotypes with diverse combination of characteristics associated with resistance to sorghum midge can be used in breeding programs to broaden the genetic base and increase the levels of resistance to this insect. This revised version was published online in August 2006 with corrections to the Cover Date.     

Isolation and characterization of resistance gene analogs (RGAs) from sorghum (Sorghum bicolor L. Moench)

Degenerate primers designed based on known resistant genes (R-genes) and resistance gene analogs (RGAs) were used in combinations to elucidate RGAs from Sorghum bicolor, cultivar M 35-1. Most of the previously tried primer combinations resulted in amplicons of expected 500–600 bp sizes in sorghum along with few novel combinations. Restriction analysis of PCR amplicons of expected size revealed a group of fragments present in a single band indicating the heterogeneous nature of the amplicon. Many of these were cloned and some were considered for analysis. The nucleotide sequence of different cloned fragments was done and their predicted amino acid sequences compared to each other and to the amino acid sequences of known R-genes revealed significant sequence similarity. A cluster analysis based on neighbor-joining (N-J) method was carried out using sorghum RGAs (SRGAs) together with several analogous known R-genes resulting in two major groups; cluster-I comprising only SRGAs and cluster-II comprised of known R-gene sequences along with three SRGAs. Further analysis clearly indicated similarity of SRGAs in overall sense with already known ones from other crop plants. These sequences can be used as guidelines to detect, map and eventually isolate numerous R-genes in sorghum.     

Genotypic resistance in sorghum to head bug,Calocoris angustatus Lethiery

Summary Sorghum head bug, Calocoris angustatus Leth., is an important pest of grain sorghum. We screened nearly 15000 germplasm accessions for resistance to this pest between 1980 and 1990 under natural and headcage conditions. Data were recorded on bug numbers, grain damage (1 = highly resistant the 5 = highly susceptible), and seed germination. Under natural conditions, 34 genotypes suffered moderate levels of grain damage (damage rating (DR) 1.7 to 2.9) compared with a DR of 4.0 to 4.6 in the susceptible controls CSH 1, CSH 5 and CSH 9. IS 17610, IS 17645, IS 21444, IS 19948, IS 25069 and IS 19949 suffered a DR of less than three, and harbored less than 150 bugs/panicle compared with a DR of 4.3 to 4.7, and 248 to 353 bugs/panicle in the susceptible controls CSH 1, CSH 5 and CSH 9 when infested under headcage with 5 pairs of bugs/panicle. IS 18274, IS 20664, IS 20059, IS 25069, and IS 19951 had 150 to 300 bugs/panicle but suffered moderate levels of grain damage (DR less than 3), while the reverse was true in case of IS 8064, IS 19455, IS 19955, IS 20024, IS 20740, IS 23627, IS 2761, and IS 9692. During the 1989 rainy season, IS 14108, IS 17610, IS 17618, IS 17645, IS 19949, IS 19950, IS 19957, IS 20068, IS 25760, IS 27452, IS 27477 and IS 27329 suffered moderate levels of grain damage when infested with 5 and 10 pairs of bugs/panicle, and recorded more than 80% seed germination compared with a DR of 3.9 to 5.0, and seed germination of 15–18% in the susceptible controls CSH 1, CSH 5 and CSH 9. There is a considerable diversity in the genotypes resistant to head bugs, and attempts should be made to transfer the resistance into agronomically acceptable cultivars.     

A genetic study of popping quality in Sorghum (Sorghum bicolor (L.) Moench)

Summary Two crosses of sorghum, Sorghum bicolor (L.) Moench (IS 1054 × ICSV-1, and IS 5604 × IS 1054) were evaluated in parental, F1, F2, and backcross generations for the variation in their popping quality as measured by pop volume (ml). Dominance was in the direction of low pop volume. Dominance and additive gene effects, in that order, governed most of the variation, while significant dominance x dominance type of interaction effects could also be detected. There was no evidence for higher order gene interactions.Approved as Journal Article 630 by the International Crops Research Institute for the Semi-Arid Tropics (ICRISAT), Patancheru, P.O. 502 324, Andhra Pradesh, India.     

Genetic analysis of grain mould resistance in coloured sorghum genotypes

Grain moulds are a major constraint to sorghum production and to adoption of improved cultivars in many tropical areas. Information on the inheritance of grain mould reaction is required to facilitate breeding of resistant cultivars. The genetic control of grain mould reaction was studied in 7 crosses of 2 resistant sorghum genotypes. P₁, P₂, F₁, F₂, BC₁ and BC₂ families of each cross were evaluated under sprinkler irrigation for field grade and threshed grade scores and subjected to generation mean analysis. Frequency distributions for grain mould reaction were derived and F₂ and BC₁ segregation ratios were calculated. Grain mould reaction in crosses of coloured grain sorghum was generally controlled by two or three major genes. Resistance to grain moulds was dominant. Significant additive gene effects were also found in all cross/season combinations. Significant dominance effects of similar magnitude to additive effects were also observed in five out of ten cross/season combinations. Gene interactions varied according to the parents with both resistant and susceptible parents contributing major genes. Choice of parents with complementary resistance genes and mechanisms of resistance will be critical to the success of resistance breeding. This revised version was published online in July 2006 with corrections to the Cover Date.     

Genetic analysis of biotype I greenbug resistance in sorghum

The greenbug has been a major insect pest of sorghum since 1968. Although sources of genetic resistance have been identified to combat this pest, new and virulent biotypes have successfully overcome these resistance genes. KS 97 was developed and released by the Kansas Agricultural Experiment Station as a new germplasm source of biotype I greenbug resistance in sorghum. The objectives of this study were to evaluate combining ability effects for greenbug resistance in KS 97 and to determine the number of genes responsible for this trait. Six inbred lines, including KS 97 and greenbug resistant and susceptible checks, were intercrossed using a Design-II mating scheme to produce nine F1 hybrids. Responses of seedlings of parent lines and hybrids to biotype I greenbug were evaluated in replicated growth chamber experiments. The results of these studies indicated significant effects of general combining ability (GCA) and specific combining ability (SCA). Greenbug resistance derived from KS 97 was found to be incompletely dominant, and the GCA effect for resistance associated with KS 97 was superior to that associated with PI550610, the resistant check. Segregation studies to determine the number of genes responsible for greenbug resistance in KS 97 were conducted in BC1F1 populations. KS 97 was introgressed into three greenbug-susceptible genetic backgrounds. Segregation analysis indicated a consistent 1:3 (resistant:susceptible) segregation ratio for greenbug resistance across populations. The simplest explanation for these results is that two dominant genes requiring complementary gene action control resistance. This revised version was published online in August 2006 with corrections to the Cover Date.     

Environmental factors influence the expression of resistance to sorghum midge, Stenodiplosis sorghicola

Host plant resistance is an effective means of controlling sorghum midge (Stenodiplosis sorghicola). We studied the influence of environmental factors on expression of resistance to sorghum midge in three midge-resistant and two midge-susceptible genotypes. Midge-resistant lines AF 28, ICSV 197, and TAM 2566 suffered 8.8 to 17.3% damage across seven so wings compared to 25.6%damage in ICSV 112, and 69.4% damage in CSH 5. Susceptibility of the midge-resistant lines (AF 28, ICSV 197, and TAM 2566) decreased with an increase in open pan evaporation, maximum and minimum temperatures, and solar radiation; while the midge-susceptible lines (ICSV 112 and CSH 5) showed a poor interaction with these factors. Midge damage in ICSV 197 showed a negative correlation with minimum temperature and relative humidity and positive correlation with sunshine hours,while the reverse was true for CSH 5. Grain growth rate between 0 and 3 days after anthesis was lower in crops sown on 1st October, when AF 28 and ICSV 197 suffered maximum midge damage. Maximum and minimum temperatures and maximum relative humidity influenced the moisture content of the grain, grain growth rate, and sorghum midge damage. There was considerable variation in genotype × environment interaction for expression of resistance to sorghum midge,and the implications of these results have been discussed in relation to development of sorghum cultivars with resistance to this insect. This revised version was published online in August 2006 with corrections to the Cover Date.     

A single recessive gene for stem sweetness in sorghum

Summary The inheritance of stemsweetness in sorghum has followed a qualitative pattern, non-sweet being monogenically dominant over sweet. Hence sweetness was controlled by a single recessive gene. A maternal effect was not involved for the expression of sweetness.     

Geographical patterns of morphological variation in sorghum (Sorghum bicolor (L.) Moench) germplasm from Ethiopia and Eritrea: Quantitative characters

A total of 415 sorghum (Sorghum bicolor (L.) Moench) accessions representing different regions of Ethiopia, Eritrea and a group of introduced lines were evaluated for 15 quantitative characters to determine the extent and geographical pattern of morphological variation. The extent of variation was highly pronounced for agronomically important characters for sorghum. These characters included plant height, days for 50% flowering, peduncle exsertion, panicle length and width, number and length of primary branches per panicle and thousand seed weight. Significant regional variation was also observed for most of the characters. The results implied that environmental factors such as altitude, rainfall, temperature and growing period are important in regional variation. Mean for plant height and for days for 50% flowering showed clinal variation along the gradients of rainfall pattern and growing period in Ethiopia. Moreover, there were significant positive correlation coefficients between most of the characters. This included the correlation between agronomic characters of primary interest in sorghum breeding such as plant height and days for 50% flowering and also between various characters and the altitude of the collection sites. The implications of the results in plant breeding, germplasm collection and conservation as well as the probable sources of the wide range of variation are discussed. This revised version was published online in July 2006 with corrections to the Cover Date.     

Cytoplasmic male-sterility and source of pollen influence the expression of resistance to sorghum midge, Stenodiplosis sorghicola

Sorghum midge, Stenodiplosis (Contarinia) sorghicola (Coquillett), is an important pest of grain sorghum, and host plant resistance is an important aspect of control of this pest. This research investigated how cytoplasmic male-sterility and source of pollen influence the expression of resistance to sorghum midge. Sorghum midge emergence was significantly lower in panicles of midge-resistant and midge-susceptible cytoplasmic male-sterile lines when pollinated with AF 28 - a midge-resistant restorer line, than those pollinated with Swarna - a midge susceptible restorer line, indicating the presence of xenia effects. Maintainer lines (B-lines) of midge-resistant parents had significantly lower numbers of eggs and larvae than the B-lines of midge-susceptible parents. Male-sterile lines of the both midge-resistant and midge-susceptible lines were equally susceptible, indicating that resistance to sorghum midge is influenced by factors in the cytoplasm of the B-line. These findings will have an important bearing on the production of hybrids with resistance to insects. This revised version was published online in August 2006 with corrections to the Cover Date.     

Evaluation of greenhouse inoculation techniques to screen sorghum for resistance to downy mildew

Summary Six inoculation techniques were compared for the artificial promotion of downy mildew (Peronosclerospora sorghi) in sorghum. These were (1) sprouted seeds incubated between sporulating infected leaves, (2) sprouted seeds depped in conidial suspension, (3) sprouted seeds sprayed with conidial suspension, (4) seedlings at plumule stage inoculated with drops of a conidial suspension, (5) seedlings at plumule stage sprayed with a conidial suspension, and (6) seedling showered with conidia falling from infected leaves. Seedlings at the one-leaf stage sprayed with a conidial suspension (6 × 10⁵ ml^-1) showed the highest systemic infection (100%) in the susceptible lines IS 643 and IS 18433. This technique is effective, repeatable, and allows the deposition of a conidial suspension as a fine mist on the entire seedling surface. In the greenhouse, the technique was used to test the downy mildew reaction of genotypes previously reported as resistant (< 5% incidence) in 3–4 years of field screenings. Of the 61 genotypes tested, 21 were free from downy mildew, 14 had less than 5% incidence, and the rest showed variable susceptible reactions. Therefore, the technique can be reliably and effectively used in the greenhouse to detect disease escapes and to indentify resistance.     

Effect of cytoplasmic male-sterility in sorghum on host plant interaction with sorghum midge,Contarinia sorghicola

Summary Sorghum midge, Contarinia sorghicola Coq. (Diptera: Cecidomyiidae) is one of the most important pests of grain sorghum worldwide. We studied the reaction of midge-resistant and midge-susceptible genic-cytoplasmic male-sterile (A-lines) and their maintainers (B-lines), and the effect of resistant and susceptible restorers on sorghum midge. Midge damage and adult emergence were significantly lower on the B-lines of midge-resistant genotypes (PM 7061 and PM 7068) than their corresponding A-lines, while the reverse was true for the midge-susceptible genotypes (296A and ICSA 42). Differences in midge damage and the number of midges emerged were not significant between the midge-resistant and midge-susceptible A-lines when infested without pollination (except midge emergence on PM 7061A). Pollination with a midge-resistant restorer (DJ6541) reduced midge emergence significantly in one of two seasons. Source of pollen did not influence midge emergence on the highly-resistant A-line, PM 7061A. The implications of these observations in the development of midge-resistant hybrids were discussed.     

Diallel analysis of sooty stripe resistance in sorghum

Sooty stripe [Ramulispora sorghi (Ellis and Everhart) Olive and Lefebre] is a widespread foliar disease of sorghum [Sorghum bicolor (L.) Moench] in West Africa, responsible for grain yield losses up to 46%. We studied the inheritance of sooty stripe resistance in a 9 × 9 sorghum F₂-population diallel grown together with parent lines and checks in1996 under natural disease pressure at two locations in Mali. The percentage of infected leaf area was determined twice over a two-week interval during the season. At the second evaluation, the mean sooty stripe severity amounted to 13% infected leaf area at Samanko and 12% at Cinzana. The frequency distribution of the entries was approximately normal for the mean disease severity, averaged across assessment dates and locations, pointing to the involvement of multiple genes. With the data combined across the two locations, genetic differences among lines and among F₂ populations were highly significant. Genotype × location interaction variances were also significant but much smaller than the genetic variances. Broad-sense heritability estimates were 0.92 for lines and 0.94 for the F₂ populations, for the mean percentage infected leaf area across the two assessment dates. General combining ability effects (GCA) determined most of the differences among the F₂ populations. Specific combining ability effects (SCA), and the interactions of GCA or SCA with locations were also significant but less important. Line performance per se was highly correlated with GCA. Because of the high heritability and predominance of additive effects, prospects are good for the genetic improvement of resistance to sooty stripe in sorghum in Mali, using simple pedigree or recurrent selection procedures. This revised version was published online in August 2006 with corrections to the Cover Date.     

Inheritance and field performance of transgenic Korean Bt rice lines resistant to rice yellow stem borer

Transgenic Korean rice plants containing the cry1Ab gene were developed for resistance against yellow stem borer (Scirpophaga incertulas, YSB). More than 100 independent transgenic lines from three Korean varieties (P-I, P-II and P-III) were generated. The amount of Cry1Ab in transgenic T₀ plants was as high as 2.88% of total soluble proteins. These levels were sufficient to cause 100% mortality of YSB larvae. The majority of T₁ transgenic lines originated from the varieties P-I and P-II followed a Mendelian fashion of segregation. Deviation from the expected segregation ratio was observed in a small number of the transgenic lines of P-I and P-II origins. However, this deviation was primarily observed in the P-III originated lines. Segregation analysis of the T₁ generation indicated that 1–3 copies of the cry1Ab gene were integrated into the genome of the majority of the transgenic lines originating from varieties P-I and P-II. Stunted and semi-fertile mutants were observed in some transgenic lines. These aberrations were either independent or closely linked to the introduced cry1Ab gene loci in different transgenic lines. Reduction in GUS expression levels and loss of toxicity against YSB larvae were found in some transgenic lines. The transgenic T₃ and T₄ lines causing 100% mortality of third instar YSB larvae in the lab were completely protected in the field. Analysis of important yield components on nine selected transgenic lines indicated that stem length, panicle length, grain number per panicle, and seed setting rates were reduced in transgenic plants compared to those in non-transgenic parental rice lines. Number of panicles per cluster, however, was significantly higher in transgenic plants. The numerical value of the average yield was in general greater in the controls than in all the transgenic lines, indicating some ‘yield drag’. Since some selected lines were highly resistant to the YSB with good yielding potential, they offer effective potential for use in insect resistance management programs.     

Heritability estimates of variation for NaCl tolerance in Sorghum bicolor (L.) Moench seedlings

Summary Data on root lengths of two-week-old seedlings of 51 Sorghum bicolor (L.) Moench accessions germinated in 50, 100, 150, 175, and 200 mM NaCl solutions were used to estimate broad sense heritabilities of variation for salinity tolerance.Estimates of broad sense heritabilities ranged from 0.38 to 0.73, the highest being in 50 mM NaCl, suggesting that there is potential for improving salinity tolerance in the species through selection and breeding.     

Genetic analysis of grain mold resistance in white seed sorghum genotypes

Grain molds in rainy season sorghums can cause poor grain quality resulting in economic losses. Grain molds are a major constraint to the sorghum production and for adoption of the improved cultivars. A complex of fungi causes grain mold. Information on genetics of grain mold resistance and mechanisms is required to facilitate the breeding of durable resistant cultivars. A genetic study was conducted using one white susceptible, three white resistant/tolerant sources, and one colored resistant source in the crossing programme to obtain four crosses. P₁, P₂, F₁, BC₁, and BC₂, and F₂ families of each cross were evaluated for the field grade and threshed grade scores, under sprinkler irrigation. Generation mean analyses and frequency distribution studies were carried out. The frequency distribution studies showed that grain mold resistance in the white-grained resistance sources was polygenic. The additive gene action and additive × additive gene interaction were significant in all the crosses. Simple recurrent selection or backcrossing should accumulate the genes for resistance. Epistasis gene interactions were observed in colored resistance × white resistance cross. Gene interaction was influenced by pronounced G × E. Pooled analysis showed that environment × additive gene interaction and environment × dominant gene interaction were significant. The complex genetics of mold resistance is due to the presence of different mechanisms of inheritance from various sources. Evaluation of segregating population for resistance and selection for stable derivatives in advanced generations in different environments will be effective.     

QTL analysis and mapping of pre-harvest sprouting resistance in Sorghum

One of the most important agronomic problems in the production of sorghum [Sorghum bicolor (L.) Moench] in humid climates is pre-harvest sprouting (PHS). A molecular linkage map was developed using 112molecular markers in an F₂ mapping population derived from a cross between IS 9530 (high resistance to PHS) and Redland B2 (susceptible to PHS). Two year phenotypic data was obtained. By means of interval mapping analysis, two significant QTL were detected in two different linkage groups with LOD scores of 8.77and 4.39. Each of these two QTL individually explained approximately 53% of the phenotypic variance, but together, in a two-QTL model, they explained 83% of the phenotypic variance with a LOD score of 12.37.These results were corroborated by a one-way ANOVA in which the four flanking markers of the most likely QTL positions displayed highly significant values in theF-test, and significant variation in trait expression was associated with marker genotypic classes. The four markers with highest effect in the one-way ANOVA were also detected in the second year replication of the F₂ population, and significant genotype × environment interactions was observed. The putative relationship between PHS resistance in sorghum and the maize Vp1 gene is also discussed. This revised version was published online in July 2006 with corrections to the Cover Date.     

Genetic divergence among a non-restorer collection of sorghum (Sorghum bicolor (L.) Moench) and its relationship with heterosis

Summary A genetic diversity analysis in a collection of 171 non-restorer lines of sorghum (Sorghum bicolor (L.) Moench) using D² technique and canonical variate analysis indicated that considerable variation in grain yield has been added to the collection by the addition of lines derived from random mating populations. The efficiency of D² and canonical variate techniques in distinguishing extremely diverse genotypes was confirmed. However, the two techniques showed weak correspondence in their clusters. The F₁ hybrids of 15 diverse lines exhibited no relationship between heterosis or per se performance of crosses and diversity in their parents. Therefore, traditional plant breeding methods are being advocated.Approved by ICRISAT as Journal Article no. 435.     

Pattern analysis of genotype × environment interaction for striga resistance and grain yield in African sorghum trials

The parasitic weed Striga hermonthica (Del.) Benth. seriously limits sorghum [Sorghum bicolor (L.) Moench] production in Sub-Saharan Africa. As an outbreeder, S. hermonthica is highly variable with an extraordinary capacity to adapt to different hosts and environments, thereby complicating resistance breeding. To study genotype x environment (G x E) interaction for striga resistance and grain yield, nine sorghum lines, 36 F₂ populations and five local checks were grown under striga infestation at two locations in both Mali and Kenya. Mean squares due to genotypes and G x E interaction were highly significant for both sorghum grain yield and area under striga severity progress curve(ASVPC, a measure of striga emergence and vigor throughout the season). For grain yield, the entry x location-within-country interaction explained most of the total G x E while for ASVPC, entry x country and entry x location-within-country interactions were equally important. Pattern analysis (classification and ordination techniques) was applied to the environment-standardized matrix of entry x environment means. The classification clearly distinguished Malian from Kenyan locations for ASVPC, but not for grain yield. Performance plots for different entry groups showed differing patterns of adaptation. The ordination biplot underlined the importance of entry x country interaction for ASVPC. The F₂ derived from the cross of the striga-resistant line Framida with the striga-tolerant cultivar Seredo was the superior entry for both grain yield and ASVPC, underlining the importance of combining resistance with tolerance in striga resistance breeding. The observed entry x country interaction for ASVPC may be due to the entries' different reactions to climatic conditions and putative differences in striga virulence in Mali and Kenya. This revised version was published online in August 2006 with corrections to the Cover Date.