A new aspect of genetic diversity of Indonesian oil palm (Elaeis guineensis Jacq.) revealed by isoenzyme and AFLP markers and its consequences for breeding

Oil palm (Elaeis guineensis Jacq.) plays an important economic role in some countries of Southeast Asia like Indonesia, which is the world’s second producer of palm and palm kernel oil. The quality improvement of planting material needs a better understanding of the genetic relationships between genotypes from different populations used in the breeding programmes. In this study, 48 parents, representative of four populations used in Indonesian Oil Palm Research Institute (IOPRI) breeding programmes, were analysed with five selected AFLP primer pairs and four isoenzymatic systems. One hundred and fifty eight scorable band levels were generated of which 96 (61%) were polymorphic. AFLP allowed us to identify off–type descendants which were excluded from analysis. The use of unbiased Rogers distance clearly separated the four studied populations. The Neighbor-Joining method re-groups two African populations which are known as originating from different regions. Nevertheless, the variability revealed is in accordance with oil palm breeders’ knowledge. The results obtained with AFLP showed that the crosses among the African sub-population, which is excluded in oil palm reciprocal recurrent selection (RRS) breeding programmes, may be more interesting than the crosses between the African and the Deli populations. Received: 9 August 1999 / Accepted: 28 February 2000     

Multiple locus genome-wide association studies for important economic traits of oil palm

Palm oil has a balanced fatty acid composition and has no trans fat. As a result, its use in food has increased as food-labeling laws have changed to specify trans fat content. Increasing oil production is the main goal in oil palm breeding. Genetic mapping and genomic studies in palm trees are necessary to understand the genetic architecture of economic traits of importance for palm oil production. To help achieve this, we sampled 422 oil palms from MPOB (Malaysian Palm Oil Board)­Angola germplasm collection and measured 13 economic traits from these palms. Multi-locus genome-wide association studies (GWAS) were conducted using least absolute shrinkage and selection operator (LASSO) and genome-wide efficient mixed model analysis. We identified 19 quantitative trait loci (QTLs) for 8 traits. Of these, four Angola-specific QTLs associated with bunch components were detected on chromosomes 4, 8, and 11. These QTLs are potentially useful for introgression of desirable genes from the Angola palms to advanced breeding populations for improvement of bunch and oil yield traits. The majority of the QTLs were detected by LASSO-A, in which the p values of individual markers were calculated based on bootstrapped standard errors. Many of the detected QTLs are nearby known QTLs detected from linkage studies reported by other research groups. We also conducted genomic selection (GS) for the 13 traits and concluded that GS can be an effective tool for oil palm breeding. This is the first GWAS and GS study conducted on oil palm germplasm from Angola, and the results can be very useful in oil palm genetic studies and breeding.     

油棕鲜果穗产量构成因素的相关性和回归分析

为明确油棕鲜果穗产量构成因素之间的关系,利用相关性、回归和通径等分析方法,对58株油棕单株鲜果穗产量与其它性状相关性进行了研究。结果表明:油棕鲜果穗产量与鲜果穗重和每株果穗数的相关性最为密切。另外,在油棕高产品种选育过程中,应着重对平均果穗重、每株果穗数、果实大小和含油量等性状进行选择,并综合权衡这些产量性状之间的相互作用。产量性状变异和相关性分析表明,平均种壳厚度变异程度最大(88.34%),平均果长变异最小(11.99%),鲜果穗产量与鲜果穗重、每株果穗数、平均果重呈极显著正相关关系(0.882**,0.714**,0.439**),与平均果长呈显著正相关关系(0.334*)。多元线性回归和通径分析结果表明,通过比较多元线性回归方程的相关参数,建立了可实用的最优多元线性回归方程x_1=-281.511+2.787x_4+3.455x_5-2.672x_6+0.411x_7+6.690x_8+3.664x_9,平均鲜果穗重和每株果穗数对单株鲜果穗产量的直接作用最大(0.670,0.439),其它性状对单株产量的直接作用较小,平均果重和平均果长对单株鲜果穗产量的间接作用最大(0.335,0.322)。该研究结果为油棕高产品种选育提供了依据。     

Mapping QTL for leaf area in oil palm using genotyping by sequencing

Leaf area is an important parameter in oil palm breeding as it is positively correlated with oil yield. However, measurement of leaf area is tedious and also destructive. In the present study, a breeding population with 145 palms derived from a cross between Deli Dura and Avros Pisifera was used to construct a high-density linkage map and identify quantitative trait loci (QTL) for leaf area in oil palm. Using genotyping by sequencing, a linkage map containing 2413 SNPs was constructed. The total length of the linkage map was 1161.89 cM, with an average marker spacing of 0.48 cM. Based on the continuous phenotyping of leaf area from 2010 to 2015, two suggestive QTL for leaf area were mapped on chromosomes (Chr) 3 and 9. The allelic effects of the QTL associated with leaf area in the mapping population were estimated by linear regression using ordinary least squares method. The QTL on Chr 9 explained 13.3% of phenotypic variation for leaf area. A candidate gene, ARC5, within the narrow interval of QTL on Chr 9 was identified. The gene was significantly higher expressed in leaf than root and fruit of oil palm. This high-quality and SNP-based map supplies a base to fine map QTL for agronomic traits in oil palm, and the markers closely linked to the stable QTL may be used in marker-assisted selection in oil palm breeding.     

Possible sources of genetic resistance in oil palm (<Emphasis Type="Italic">Elaeis guineensis</Emphasis> Jacq.) to basal stem rot caused by <Emphasis Type="Italic">Ganoderma boninense</Emphasis> – prospects for future breeding

Oil palm estates in southeast Asia suffer from substantial losses due to basal stem rot caused by Ganoderma boninense. Field observations have been carried out in North Sumatra, Indonesia, on a series of planting materials of known origin. Differences in susceptibility to the disease have been detected within the two Elaeis species, guineensis and oleifera. Within Elaeis guineensis, material of Deli origin is highly susceptible compared to material of African origin. It is also possible to detect differences in reaction between parents and between crosses within a given origin. The variability of resistance to basal stem rot within the same cross is also illustrated by the diverse responses of clones derived from palms of the same origin. The prospects opened up by these results are discussed, and the importance of performing an early selection test is highlighted.     

Microsatellite Analysis of Oil Palms and their Progenies Bred in Papua New Guinea

Our aim was to assess parental and population genetic variability in a partially nested set of breeding populations from the breeding program at Dami Research Station (West New Britain, PNG). Twenty microsatellite markers were tested for their ability to characterize genetic variation in oil palm populations bred at Dami Research Station. One hundred and twenty six individuals, including 100 F₁ tenera hybrids of Dami Deli crossed with either AVROS, Ghana or AVROS/Ghana breeding lines were analysed. Eighteen of these markers were polymorphic within and among populations, amplifying 103 alleles in oil palm. Three individuals of other palm species (Cocos nucifera and two Phoenix spp.) were included as outliers. With these markers we have the power to distinguish individual palms, hence we conclude that they will facilitate association of markers with important phenotypic traits to streamline future breeding and selection.     

Mapping of quantitative trait loci (QTLs) for oil yield using SSRs and gene-based markers in African oil palm (<Emphasis Type="Italic">Elaeis guineensis</Emphasis> Jacq.)

The identification of quantitative trait loci (QTLs) based on anchor markers, especially candidate genes that control a trait of interest, has been noted to increase the power of QTL detection. Since these markers can be scored as co-dominant data, they are also valuable for comparing and integrating the QTL linkage maps from diverse mapping populations. To estimate the position and effects of QTLs linked to oil yield traits in African oil palm, co-dominant microsatellites (SSR) and candidate gene-based sequence polymorphisms were applied to construct a linkage map for a progeny showing large differences in oil yield components. The progeny was genotyped for 97 SSR markers, 93 gene-linked markers, and 12 non-gene-linked SNP markers. From these, 190 segregating loci could be arranged into 31 linkage groups while 12 markers remained unmapped. Using the single marker linkage, interval mapping and multiple QTL methods, 16 putative QTLs on seven linkage groups affecting important oil yield related traits such as fresh fruit bunch yield (FFB), ratio of oil per fruit (OF), oil per bunch (OB), fruit per bunch (FB) and wet mesocarp per fruit (WMF) could be identified in the segregating population with estimated values for explained variance ranging from 12.4 % to 54.5 %. Markers designed from some candidate genes involved in lipid biosynthesis were found to be mapped near significant QTLs for various economic yield traits. Associations between QTLs and potential candidate genes are discussed.     

Long-term genomic selection for heterosis without dominance in multiplicative traits: case study of bunch production in oil palm

Background

To study the potential of genomic selection for heterosis resulting from multiplicative interactions between additive and antagonistic components, we focused on oil palm, where bunch production is the product of bunch weight and bunch number. We simulated two realistic breeding populations and compared current reciprocal recurrent selection (RRS) with reciprocal recurrent genomic selection (RRGS) over four generations. All breeding strategies aimed at selecting the best individuals in parental populations to increase bunch production in hybrids. For RRGS, we obtained the parental genomic estimated breeding values using GBLUP with hybrid phenotypes as data records and population specific allele models. We studied the effects of four RRGS parameters on selection response and genetic parameters: (1) the molecular data used to calibrate the GS model: in RRGS_PAR, we used parental genotypes and in RRGS_HYB we also used hybrid genotypes; (2) frequency of progeny tests (model calibration); (3) number of candidates and (4) number of genotyped hybrids in RRGS_HYB.

Results

We concluded that RRGS could increase the annual selection response compared to RRS by decreasing the generation interval and by increasing the selection intensity. With 1700 genotyped hybrids, calibration every four generations and 300 candidates per generation and population, selection response of RRGS_HYB was 71.8 % higher than RRS. RRGS_PAR with calibration every two generations and 300 candidates was a relevant alternative, as a good compromise between the annual response, risk around the expected response, increased inbreeding and cost. RRGS required inbreeding management because of a higher annual increase in inbreeding than RRS.

Conclusions

RRGS appeared as a valuable method to achieve a long-term increase in the performance for a trait showing heterosis due to the multiplicative interaction between additive and negatively correlated components, such as oil palm bunch production.     

Character interrelationships and path coefficient analysis for oil yield in the oil palm

Path coefficient analysis of eight yield components correlated with yield in 22 oil palm progenies showed that number of bunches per palm had the largest direct effect on oil yield followed single bunch weight and percent oil-to-mesocarp ratio.     

Genetic linkage map of a high yielding FELDA deli×yangambi oil palm cross

Enroute to mapping QTLs for yield components in oil palm, we constructed the linkage map of a FELDA high yielding oil palm (Elaeis guineensis), hybrid cross. The parents of the mapping population are a Deli dura and a pisifera of Yangambi origin. The cross out-yielded the average by 8-21% in four trials all of which yielded comparably to the best current commercial planting materials. The higher yield derived from a higher fruit oil content. SSR markers in the public domain - from CIRAD and MPOB, as well as some developed in FELDA - were used for the mapping, augmented by locally-designed AFLP markers. The female parent linkage map comprised 317 marker loci and the male parent map 331 loci, both in 16 linkage groups each. The number of markers per group ranged from 8-47 in the former and 12-40 in the latter. The integrated map was 2,247.5 cM long and included 479 markers and 168 anchor points. The number of markers per linkage group was 15-57, the average being 29, and the average map density 4.7 cM. The linkage groups ranged in length from 77.5 cM to 223.7 cM, with an average of 137 cM. The map is currently being validated against a closely related population and also being expanded to include yield related QTLs.     

Training genomic selection models across several breeding cycles increases genetic gain in oil palm in silico study

Genomic selection (GS) is expected to increase the rate of genetic gain in oil palm. In a GS scheme, breeding cycles with progeny tests (phenotypic selection, PS) used to calibrate the GS predictive model and for selection alternate with GS cycles, making it possible to train the GS model with aggregated data from several cycles. To evaluate this possibility, we simulated four cycles of hybrid breeding for bunch production and compared two methods of calibrating the GS model, one using aggregated data from the two most recent cycles (Tr2Gen), the other using data from the last cycle (Tr1Gen). We also compared a GS scheme with two PS cycles and two GS cycles (2PT-2noPT), and a scheme with PS every other cycle and GS otherwise (PT-noPT). We showed that Tr2Gen had a 10.7% higher genetic gain per cycle than Tr1Gen, mostly due to increased selection accuracy, particularly in across-cycle selection, despite the decreased relationship between training individuals and selection candidates. After four cycles, Tr2Gen had a 5% higher cumulative genetic gain than Tr1Gen, with a lower coefficient of variation. PT-noPT benefited more from the advantages offered by Tr2Gen than 2PT-2noPT. Over four breeding cycles, combining PT-noPT and Tr2Gen largely outperformed conventional reciprocal recurrent selection (RRS), with an increase in annual genetic gain ranging from 37.6 to 57.5%, depending on the number of GS candidates. This study confirms the advantages of GS over RRS and indicated that oil palm breeders should train GS models using all data from past breeding cycles.     

SSR markers in transcripts of genes linked to post-transcriptional and transcriptional regulatory functions during vegetative and reproductive development of Elaeis guineensis

Background  

The oil palm (Elaeis guineensis Jacq.) is a perennial monocotyledonous tropical crop species that is now the world's number one source of edible vegetable oil, and the richest dietary source of provitamin A. While new elite genotypes from traditional breeding programs provide steady yield increases, the long selection cycle (10-12 years) and the large areas required to cultivate oil palm make genetic improvement slow and labor intensive. Molecular breeding programs have the potential to make significant impacts on the rate of genetic improvement but the limited molecular resources, in particular the lack of molecular markers for agronomic traits of interest, restrict the application of molecular breeding schemes for oil palm.     

Phenological stages and degree days of oil palm crosses grown under irrigation in tropical conditions

Studies on phenology and growing degree days (GDD) of four oil palm crosses, that is, Palode, Deli × Nigeria, United Plantations and Deli × Ghana were conducted in India under tropical conditions. Observations were recorded in adult oil palms over a period of 1.5 years from visual opening of spear leaf to development of female flower till harvest. Biologische Bundesantalt, Bundessortenamt and Chemische Industrie (BBCH) General Scale was used for conducting various phenological growth stages. The GDD from development of spear leaf to maturity in the different crosses varied between 6,320.2 and 6,937.3. The degree days and duration from development of spear leaf to maturity were less in the crosses of United Plantations. The time taken for spear leaf unfolding to maturity and flower opening to maturity in the different crosses, respectively, varied from 447.9 to 485.2 days and 145.8 to 153.7 days. The study gives an insight in evaluating thermal time for achieving various phenophases in oil palm and genotypic variations of time taken in attaining the different phenophases that have been documented. Climatic and yield prediction models can also be evolved through these studies.     

Microsatellite-based high density linkage map in oil palm (Elaeis guineensis Jacq.).

A microsatellite-based high-density linkage map for oil palm (Elaeis guinensis Jacq.) was constructed from a cross between two heterozygous parents, a tenera palm from the La Mé population (LM2T) and a dura palm from the Deli population (DA10D). A set of 390 simple sequence repeat (SSR) markers was developed in oil palm from microsatellite-enriched libraries and evaluated for polymorphism along with 21 coconut SSRs. A dense and genome-wide microsatellite framework as well as saturating amplified fragments length polymorphisms (AFLPs) allowed the construction of a linkage map consisting of 255 microsatellites, 688 AFLPs and the locus of the Sh gene, which controls the presence or absence of a shell in the oil palm fruit. An AFLP marker E-Agg/M-CAA132 was mapped at 4.7 cM from the Sh locus. The 944 genetic markers were distributed on 16 linkage groups (LGs) and covered 1,743 cM. Our linkage map is the first in oil palm to have 16 independent linkage groups corresponding to the plants 16 homologous chromosome pairs. It is also the only high-density linkage map with as many microsatellite markers in an Arecaceae species and represents an important step towards quantitative trait loci analysis and physical mapping in the E. guineensis species.     

Phenology and growth adjustments of oil palm (Elaeis guineensis) to photoperiod and climate variability

Background and Aims

Oil palm flowering and fruit production show seasonal maxima whose causes are unknown. Drought periods confound these rhythms, making it difficult to analyse or predict dynamics of production. The present work aims to analyse phenological and growth responses of adult oil palms to seasonal and inter-annual climatic variability.

Methods

Two oil palm genotypes planted in a replicated design at two sites in Indonesia underwent monthly observations during 22 months in 2006–2008. Measurements included growth of vegetative and reproductive organs, morphology and phenology. Drought was estimated from climatic water balance (rainfall – potential evapotranspiration) and simulated fraction of transpirable soil water. Production history of the same plants for 2001–2005 was used for inter-annual analyses.

Key Results

Drought was absent at the equatorial Kandista site (0°55′N) but the Batu Mulia site (3°12′S) had a dry season with variable severity. Vegetative growth and leaf appearance rate fluctuated with drought level. Yield of fruit, a function of the number of female inflorescences produced, was negatively correlated with photoperiod at Kandista. Dual annual maxima were observed supporting a recent theory of circadian control. The photoperiod-sensitive phases were estimated at 9 (or 9 + 12 × n) months before bunch maturity for a given phytomer. The main sensitive phase for drought effects was estimated at 29 months before bunch maturity, presumably associated with inflorescence sex determination.

Conclusion

It is assumed that seasonal peaks of flowering in oil palm are controlled even near the equator by photoperiod response within a phytomer. These patterns are confounded with drought effects that affect flowering (yield) with long time-lag. Resulting dynamics are complex, but if the present results are confirmed it will be possible to predict them with models.     

棉花亲本选配模式的研究

根据棉花的数量性状,利用聚类分析方法将１７个亲本分为４类．根据棉花的综合性状和性状距离,给出了两套亲本选配方案．试验结果表明,这些方案能提高育种工作中亲本选配的效率．     

Breeding for reduced seed coat fuzz in Upland cotton (Gossypium hirsutum)

The inheritance of seed coat fuzz was studied in two half diallel sets of crosses of Upland cotton. One with F₄ selections from an inter-varietal cross showed a significant level of non-additive variance attributable to dominance and non-allelic interaction. In the other, using inbred varieties of diverse origin, the genetic control of seed fuzz was adequately accounted for by an additive-dominance model with no interaction. Genotypic correlations between seed coat fuzz, yield and lint quality characters, calculated for both diallel sets and for two other groups of breeding material, showed good agreement within each experiment between parents and hybrids or between parents and progenies but no consistent pattern between experiments. The results serve to emphasize the risks in extrapolating correlations from one group of breeding material to another. A useful level of reduced fuzz has been obtained in selections from the AH breeding programme and the genetical investigations indicate that a further reduction may be possible, thereby leading to easier handling of seed, speedier and cheaper ginning, low levels of seed coat nep and better seed germination.     

Oil Palm Empty Fruit Bunch and Sugarcane Bagasse Enhance the Bioremediation of Soil Artificially Polluted by Crude Oil

Contamination of soil by petroleum hydrocarbons is becoming prevalent in Malaysia. Infiltration of soil contamination into groundwater poses a great threat to the ecosystem and human health. Bioremediation can occur naturally or can be enhanced with supplementation of microorganisms and fertilizers. However, fertilizers are expensive and therefore alternative nutrient-rich biomaterials are required. In this study, two organic wastes from agricultural industry (i.e., sugarcane bagasse and oil palm empty fruit bunch) were investigated for possible enhanced bioremediation of soil contaminated with Tapis crude oil. Two bacterial strains isolated and characterized previously (i.e., Pseudomonas aeruginosa UKMP-14T and Acinetobacter baumannii UKMP-12T) were used in this study. Sugarcane bagasse (5% and 15%, w/w) and oil palm empty fruit bunch (20%, w/w) were mixed with soil (500 g) spiked with Tapis crude oil (3%, v/w). The treated soils as well as controls were incubated for 20 days under controlled conditions. Sampling was carried out every four days to measure the number of bacterial colonies (CFU/g) and to determine the percentage of oil degradation by gas chromatography. The two biostimulating agents were able to maintain the soil moisture holding capacity, pH, and temperature at 38-40% volumetric moisture content (VMC), 7.0, and 29–30°C; respectively. The growth of bacteria consortium after 20 days in the treatment with sugarcane bagasse and oil palm empty fruit bunch had increased to 10.3 CFU/g and 9.5 CFU/g, respectively. The percentage of hydrocarbon degradation was higher in the soil amended with sugarcane bagasse (100%) when compared to that of oil palm empty fruit bunch (97%) after 20 days. Our results demonstrated the potential of sugarcane bagasse and oil palm empty fruit bunch as good substrates for enhanced bioremediation of soil contaminated with petroleum crude oil.     

Exploiting an oil palm EST database for the development of gene-derived SSR markers and their exploitation for assessment of genetic diversity

A total of 5,521 expressed sequence tags (ESTs) from oil palm were used to search for type and frequency of simple sequence repeat (SSR) markers. Dimeric repeat motifs appeared to be the most abundant, followed by tri-nucleotide repeats. Redundancy was eliminated in the original EST set, resulting in 145 SSRs in 136 unique ESTs (114 singletons and 22 clusters). Primers were designed for 94 (69.1%) of the unique ESTs (consisting of 14 consensus and 80 singletons). Primers for 10 EST-SSRs were developed and used to evaluate the genetic diversity of 76 accessions of oil palm originating from seven countries in Africa, and the standard Deli dura population. The average number of observed and effective alleles was 2.56 and 1.84, respectively. The EST-SSR markers were found to be polymorphic with a mean polymorphic information content value of 0.53. Genetic differentiation (F _ST) among the populations studied was 0.2492 indicating high level of genetic divergence. Moreover, the UPGMA (unweighted pair-group method with arithmetic mean) analysis revealed a strong association between genetic distance and geographic location of the populations studied. The germplasm materials exhibited higher diversity than Deli dura, indicating their potential usefulness in oil palm improvement programmes. The study also revealed that the populations from Nigeria, Congo and Cameroon showed the highest diversity among the germplasm evaluated in this study. The EST-SSRs further demonstrated their worth as a new source of polymorphic markers for phylogenetic analysis, since a high percentage of the markers showed transferability across species and palm taxa.