Research Centre MPOB Natsem 2005 OP Planting Materials

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1 Breeding Oil Palms for High Oil Yield DxP in IOI Group I. First cycle development of OPGL-derived materials C R Donough and Chia C C IOI Research Centre, Batang Melaka, Negri Sembilan, MALAYSIA Summary IOI Group (IOI) purchased Dunlop Estates in 1990, including its Research Centre and a small oil palm breeding programme. The Research Centre was re-named IOI Research Centre (IOI RC). In 2003, IOI bought Unipamol, along with its oil palm breeding programme and seed production operations. Pamol and Dunlop had been part of the Oil Palm Genetics Laboratory (OPGL) established in Just prior to the closure of OPGL in 1973, a limited breeding programme (LBP) was initiated to provide Pamol and Dunlop with materials that would allow them to produce seeds in future. The LBP materials were divided between the two companies and each planted their own set in After 10 years of evaluation, both companies independently started selection of seed parents from the trials in 1985, and soon after commenced production of DxP seeds on their own. Though the starting genetic materials were the same, the seed produced by each company turned out to be different because :- Selection criteria used to select dura parents were not the same, and Different pollen was used by each company. The dura parents from the LBP turned out to be excellent for qualities related to high oil yield, having thicker mesocarp, thinner shell, higher oil/mseocarp and bigger fruits. When combined with AVROS (Av) pisifera parents, as in Pamol seed, their DxP progeny were often the highest yielding for oil and EOE (estimated oil equivalent = oil yield + 60% kernel yield) through superior oil/bunch. The LBP pisiferas produced DxP progeny that easily surpassed the Malaysian Standard MS157, and were comparable to, or even better than, most other Malaysian seed. DxP progeny of the best LBP pisiferas matched, and surpassed even that from Av pisiferas, and were much shorter. However, their superiority was from high yield of fresh fruit bunches (FFB), rather than oil/bunch. The LBP pisifera population continues to be developed in the IOI breeding programme. Thus, for a programme that was called limited at the outset, this review clearly shows that the achievements made have been anything but limited. Since the Unipamol acquisition by IOI, the breeding and seed production operations of Pamol have been integrated into the IOI RC. A review and re-selection of all dura seed parents was conducted according to the Pamol criteria and completed in 2004, and the Av pisiferas used to produce Pamol seed were adopted in favour of LBP pisiferas. From January 2005, seed production management and marketing of seed has been centralized at IOI RC, and a single brand of seed IOI Deli x AVROS seed is now produced for sale. IOI is confident that as it continues to replant its estates with the new seed, its yields and profitability will grow higher. Keywords : IOI, DxP, oil yield, oil palm breeding, OPGL, Deli x AVROS the rights to all information contained herein. Page 1 of 26

2 Introduction IOI Group (IOI) entered the plantation industry in 1983 with the purchase of two estates in Peninsular Malaysia. In just two decades, IOI s plantation holdings have grown to reach 158,502 hectares by mid (IOI Corporation Berhad Annual Report 2004). This tremendous growth was achieved through a combination of acquisitions of existing plantations, as well as development of acquired unplanted land. Today, IOI is one of the largest integrated palm oil producers in the world, with a business that encompasses plantations, mills, refineries, and downstream specialty fats and oleochemical operations (IOI Corporation Berhad Annual Report 2004). IOI s direct involvement in oil palm research started with the acquisition of Dunlop Estates Berhad (Dunlop) in The Dunlop Research Centre at Gemencheh was re-named IOI Research Centre (IOI RC). Having been established since the 1950s, the newly-named IOI RC already had a long history of research in inland areas of Peninsular Malaysia on the three major plantation crops viz. rubber, oil palm and cocoa. At the time of acquisition, the research programme included a small oil palm breeding component inherited through Dunlop s earlier involvement in the Oil Palm Genetics Laboratory (OPGL) (Hardon, 1974). IOI would go on to develop this into a full-fledged oil palm planting materials development programme (Leong et al, 1998; Yong & Leong, 2000). In January 2003, IOI acquired the Unipamol Group from Unilever. This brought with it the extensive oil palm planting materials development programme (including seed production) of Pamol Plantations (Rosenquist, 1985, 1988; Donough & Law, 1995), and thus united under the IOI umbrella two oil palm breeding programmes that shared a common beginning in 1965 in the OPGL. In this paper, we review progress made in the breeding and seed production programmes related to the ex-opgl materials in IOI Group todate. The review will cover the following :- History of the OPGL limited breeding programme (LBP) for Dunlop and Pamol, and Development of OPGL-derived materials in the respective companies leading to the production of high-yielding DxP planting materials. The discussion will include, to varying degrees of detail, the following aspects :- Selection strategies and methods employed, Performance and characteristics of the seed produced, and Integration of the Pamol seed production programme into IOI Group. the rights to all information contained herein. Page 2 of 26

3 Oil Palm Genetics Laboratory a common beginning Both Dunlop and Pamol started their involvement in oil palm breeding in 1965 through their participation in, and equal funding of, the OPGL, a research consortium sponsored by 4 companies that also included the Guthrie group and Harrisons & Crosfield (H&C, now Golden Hope Plantations) as the other participants. The OPGL was established by the 4 companies to carry out investigations into the selection and breeding of oil palm, with a view to producing superior oil palm plantings for the members of the consortium (Hardon, 1974). A significant endeavour of the OPGL was the introduction of oil palm germplasm and breeding materials from outside Malaysia to bring new genetic diversity into the existing breeding programmes of the consortium partners in Malaysia (Chan et al, 1986). However, only one of these introductions elite tenera material from the Nigerian Institute for Oil Palm Research, NIFOR was planted in field trials at Dunlop (in 1967) and Pamol (in 1968). A major output from the OPGL was the development of allometric methods of estimating dry matter production in oil palms, and the application of these methods to calculate growth indices for use in oil palm breeding and physiology studies (Hardon et al, 1969; Corley et al, 1971; Hardon et al, 1972). These non-destructive methods continue to be used today. The OPGL Limited Breeding Programme for Pamol and Dunlop With the OPGL due to close in 1973, a limited breeding programme (LBP) was started in 1971 to provide Pamol and Dunlop with a starting nucleus of breeding materials in order to allow them to produce seeds later (Hardon, 1974). By 1974, seedlings of the following materials were ready for distribution to Pamol and Dunlop :- 17 dura (D) families (including 2 D selfs), 4 tenera (T) x T families, and 7 DxT families. The 7 DxT families were not related to the 17 D and 4 T families all the parent palms (25 D and 9 T) used in their creation were different. Since we are only considering development of the LBP materials for DxP seed production, the DxT families will be excluded from further discussion. Both companies planted their respective sets of LBP materials in late-1974, Pamol establishing theirs at Pamol Estate in Kluang, Johore (Donough & Law, 1995), and Dunlop establishing theirs at Paya Lang Estate in Batu Anam, Johore (Yong & Leong, 2000). Based on Dr Hardon s 1974 OPGL report, at least a subset of the crosses was also planted at Guthrie s genetic block (GB) 71. the rights to all information contained herein. Page 3 of 26

4 Ancestry of the LBP materials (a) the Dura The D parents used to produce the D families had well-established pedigrees (ancestries of 8 selected D families given in Appendix 1). Rosenquist (1985) noted that many of the ancestral palms are known to have good breeding values for oil yield, which gives confidence in the population as a source of seed parents. Rosenquist (1986a) noted that the D families in the LBP could be grouped as follows :- pure Ulu Remis (UR) Deli crosses, with ancestries tracing back to the first selections at Ulu Remis Estate in the 1930s; mixed Deli crosses with genes from UR, Serdang Avenue (SA), Socfin (SOC), Dumpy (Dy), and Klanang Bharu (KB); and crosses between Deli and duras that included non-deli (African) genes in their ancestry. Details of the families and their distribution between Pamol and Dunlop are given in Table 1. Table 1. Parentage, genetic origins and distribution of OPGL LBP 1 dura families to Pamol and Dunlop (selected families 2 in bold type) Pamol Dunlop Cross Female Male Genetic Code 3 Code 4 Type Parent Parent Origins 5 PK 114 D26 D self UR 609/316D UR 609/316D UR PK 121 DxD UR 447/9D UR 425/15D UR PK 116 D23 DxD UR 666/166D UR 609/316D UR PK 107 D13 DxD UR 603/1074D UR 666/166D UR PK 119 D24 D self UR 706/512D UR 706/512D KB PK 100 D17 DxD UR 435/7D UR 425/15D ¼ SA, ¾ UR PK 101 D15 DxD UR 665/1887D UR 460/10D ¼ SA, ¾ UR PK 105 D14 DxD UR 547/459D UR 609/316D ¼ SA, ¾ UR PK 113 DxD UR 460/10D UR 609/316D ¼ SA, ¾ UR PK 115 D16 DxD UR 654/499D UR 609/316D ¼ SA, ¾ UR PK 117 D21 DxD UR 460/10D UR 666/166D ¼ SA, ¾ UR PK 103 D11 DxD UR 452/1D UR 515/3018D ⅜ SA, ⅝ UR PK 106 D25 DxD UR 706/512D UR 609/316D ½ KB, ½ UR PK 118 D22 DxD UR 666/166D UR 706/512D ½ KB, ½ UR PK 108 D12 DxD UR 732/761D UR 666/166D ¼ SOC, ¾ UR PK 104 DxD UR 597/729D UR 515/3018D ¼ Dy, ¼ SA, ½ UR PK 122 D20 DxD UR 537/1167D UR 460/10D ⅛ Ybi, ¼ SA, ⅝ UR 1 Oil Palm Genetics Laboratory Limited Breeding Programme; 2 families selected as sources of dura seed parents; 3 cross codes assigned by Pamol Kluang (PK); 4 cross codes assigned by Dunlop (D), blank means cross not planted; 5 - refers to breeding population of restricted origin (BPRO) (Rosenquist, 1986b), UR=Ulu Remis, KB=Klanang Bharu, SA=Serdang Avenue, Dy=Dumpy, Ybi=Yangambi, SOC=Socfin. The 17 D families (15 crosses + 2 selfs) were created using 16 D parents (Table 2). According to Rosenquist (1988), these 1974 trials were among the very earliest trials to be planted with progenies from parents selected on the basis of growth records and physiological characters. Unfortunately, the data for the parents are not available in IOI RC. the rights to all information contained herein. Page 4 of 26

5 Table 2. Dura parents used in OPGL LBP 1 dura families and their origins (Parents of selected families 2 in bold type) Dura No. of Genetic Non-UR Parent 3 Crosses 4 Origins 5 Ancestors 6 UR 425/15D 2 UR UR 447/9D 1 UR UR 603/1074D 1 UR UR 609/316D 6 UR UR 665/1887D 1 UR UR 666/166D 5 UR UR 706/512D 3 KB 57/77, 68/25 UR 435/7D 1 ½ SA, ½ UR SA37, SA67 UR 460/10D 4 ½ SA, ½ UR SA5, SA7 UR 515/3018D 2 ½ SA, ½ UR SA3, SA5,SA27 UR 452/1D 1 ¼ SA, ¾ UR SA5, SA7 UR 547/459D 1 ½ SA, ½ UR SA5, SA7 UR 654/499D 1 ½ SA, ½ UR SA5, SA7 UR 597/729D 1 ½ Dy, ½ UR E206.3/8 UR 537/1167D 1 ¼ Ybi, ¾ UR Bulked Ybi pollen 7 UR 732/761D 1 ½ SOC, ½ UR Soc Oil Palm Genetics Laboratory Limited Breeding Programme; 2 families selected as sources of dura seed parents; 3 identitiy of parent from the Ulu Remis breeding programme used to create the LBP dura families; 4 no.of DxD crosses (including selfs)in which the parent was used; 5 refers to breeding population of restricted origin (BPRO) (Rosenquist, 1986b), UR=Ulu Remis, KB=Klanang Bharu, SA=Serdang Avenue, Dy=Dumpy, Ybi=Yangambi, SOC=Socfin; 6 ancestral palms that are not from the pure UR BPRO; 7 pollen imported from Congo in bulk by Oil Palms of Malaya in the rights to all information contained herein. Page 5 of 26

6 Ancestry of the LBP materials m (b) Tenera The 4 T families consisted of 2 URT x Binga T families, and 2 URT x Brabanta T families. In past reports by Yong and Leong (1998, 2000), the latter were called URT x Pindi. This was apparently a mistake in notation that went back a long way (Dr R H V Corley, personal communication). The URT parents that were used in these 2 sets of TxT families could be divided into 2 groups :- pure Yangambi origin (URTYb), and Deli-Yangambi origin (URTDeYb)(25-50% Yangambi genes) (ancestry given by Yong & Leong (2000)). The URTDeYb parents were crossed with Brabanta T pollen. The resulting progeny thus had 50-75% Deli genes, and so these crosses were not considered as sources of pisifera (P) parents for use with Deli D parents, as the resulting DxP hybrid progeny would be inbred. As such, these crosses are excluded from further discussion in this review. The URTYb parents were crossed with Binga T pollen. Yong and Leong (1998, 2000) called the resulting crosses IBY (for IOI Binga-Yangambi ), to denote that the origin of the Binga parents was actually Yangambi. Their ancestries are given in Appendix 2. Yong and Leong (2000) erroneously called one of these crosses coded D1 at Dunlop and PK112 at Pamol a Pindi-Yangambi ; the male parent, BG1032/III T (see Appendix 2), was from Binga, not Pindi as reported by Yong and Leong (loc cit). The URTYb parents trace their ancestry back via the Ivory Coast (Pobé and La Mé) to the original plantings at the Institut National pour l Étude Agronomique du Congo Belge (INEAC) in the 1920s (Rosenquist, 1986b). Yangambi materials from the INEAC station were later introduced by the IRHO (Institut pour Recherche sur les Huiles et Oléagineux, now CIRAD-CP or Centre de Coopération internationale en Recherche Agronomique pou le Dévelopment-Culture Pérennes) to their stations in Ivory Coast in the late 1940s and early 1950s (Corley & Tinker, 2003). They were probably imported as seed by Guthrie s Chemara Research Station from the Ivory Coast (Dr R H V Corley, personal communication). The two Binga T parents BG142/XXI T and BG1032/III T were imported as pollen by OPGL (Dr R H V Corley, personal communication), probably from the INEAC substation at Binga. Both Binga parents are of mainly Yangambi origin (see Appendix 2), with ancestries tracing back to descendants of the famous Djongo (Dj, the best ) palm planted at the INEAC station. BG142/XXI is pure African, with ½ Dj genes, and the other parent is from Likete (see Appendix 2). No information is available for the Likete parent. BG1032/III T had a more mixed background, with ½ Deli genes from its Socfin (Johore Labis, JL) Deli parent, and the other parent was of Isangi-Dj origins (Dr R H V Corley, personal communication). Thus, the IBY cross PK112, or D1, was ¼ Deli. This was not known at the time when selection was done in 1985, but fortuitously, this part-deli cross was not chosen as a potential source of P parents. the rights to all information contained herein. Page 6 of 26

7 First cycle selection from the LBP in IOI Group - (a) Dura parents Both Pamol and Dunlop adopted the same approach of family and individual palm selection (FIPS), as defined by Rosenquist (1990), in identifying D parents from these LBP trials in 1985 (Donough & Law, 1995; Yong & Leong, 2000). Differences arose from the following :- Data considered, in particular the vegetative measurements and growth indices, and Bias in emphasis given to different characters by those doing the selection. At Pamol, selection was done by the senior author under the guidance of Eric Rosenquist, while at Dunlop, Chiu Sheng-Bin did the selection with advice from Dr. A C Soh of Applied Agricultural Research (AAR, now re-named Applied Agricultural Resources). At the family level, selection at both sites took into account FFB yields and bunch components estimated from bunch analysis (Donough & Law, 1995; Yong & Leong, 2000; S B Chiu, personal communication). At Pamol, vegetative measurements were used to calculate dry matter production (growth) parameters and derive physiological growth indices according to the method of Corley and Breure (1981), and these were taken into account in selection (Donough & Law, 1995). Though vegetative measurements were also available at Dunlop, this was apparently not relied upon or used instead, palm form was visually assessed in the field (S B Chiu, personal communication). These differences in data consideration notwithstanding, identical D families were selected at both sites these selected families are shown in Table 4, with some data extracted from Donough and Law (1995) and Yong and Leong (2000) included for comparison. Table 4. Common Deli dura families 1 selected by Pamol and Dunlop as sources of seed parents for production of DxP hybrid seeds Pamol Dunlop Cross Female Male FFB Yield 5 Oil/bunch 6 Code 2 Code 3 Type Parent Parent Pamol Dunlop Pamol Dunlop PK 103 D11 DxD UR 4 452/1 UR 515/ PK 106 D25 DxD UR 706/512 UR 609/ PK 108 D12 DxD UR 732/761 UR 666/ PK 117 D21 DxD UR 460/10 UR 666/ PK 118 D22 DxD UR 666/166 UR 706/ PK 119 D24 D self UR 706/512 UR 706/ PK 116 D23 DxD UR 666/166 UR 609/ PK 100 D17 DxD UR 435/7 UR 425/ Mean of selected D families crosses from the Oil Palm Genetics Laboratory (OPGL) limited breeding programme for Pamol & Dunlop; 2 from Donough & Law (1995) Table 2; 3 from Yong & Leong (2000) Table 1; 4 UR=Ulu Remis, denotes that parent palm originated from the breeding programme at Ulu Remis Estate; 5 all data in kg palm -1, Pamol data from Donough & Law (1995) Table 2, Dunlop data from Yong & Leong (2000) Table 2; 6 oil/bunch in %, analysis method differed between sites Pamol used cold extraction (Blaak, 1970), Dunlop used soxhlet method ( Rao et al, 1983). the rights to all information contained herein. Page 7 of 26

8 It may be worth noting here the origins present in the ancestry of the selected D families (more details in Table 1 and Appendix 1) :- Pure UR Deli 1 family Pure KB Deli I family UR Deli + KB Deli 2 families UR Deli + SA Deli 3 families UR Deli + SOC Deli 1 family There was generally good agreement for the FFB yield data from the selected Deli progenies between the 2 sites (Table 4). However, the oil/bunch estimates were clearly different, with every corresponding result higher at the Paya Lang site, and the overall mean for the 8 selected families 18% higher there. Some of this observed bias between the two sites may be due to location effects. There was a small difference in planting density between the trials 138 palms/ha at Pamol (Donough & Law, 1995), and 148 palms/ha at Paya Lang (IOI RC, unpublished internal records) but this difference was not likely to be big enough to influence the bunch components. Soil type was similar for both sites mixed Batu Anam and Durian series with no lateritic concretions, with terrain in the trial sites flat or nearly so. The rainfall at Paya Lang ( 1,800mm a year) and its environs, however, is considerably less (by 25%) than that at the Pamol area ( 2,400mm a year) and so could be a possible factor. Parts of the bunch analysis method employed by the two research stations at that time were different. Pamol used a modification of the cold extraction scheme described by Blaak (1970), whereas Dunlop employed the more widely-used soxhlet method (Rao et al, 1983). The cold extraction method is known to give lower values for oil/mesocarp than the soxhlet method (IOI RC, unpublished data). The study of bunch analysis methods by Rao et al (1983) also showed that sampling method variations between laboratories led to large differences in results. These could be the main reasons for the observed bias in oil/bunch between Pamol and Paya Lang in Table 4. At Pamol, five families were originally selected based on their high yield of EOE (estimated oil equivalent = oil yield + 60% kernel yield, a measure that converts total products, i.e. oil + kernels, to a common price base, the 60% factor for kernel conversion being based on the long-term price differential between oil and kernels) :- Trial 74/ t/ha EOE (trial mean) Trial 74/ t/ha EOE (trial mean) PK t/ha PK t/ha PK t/ha PK t/ha PK t/ha. Three of the five families selected had palm UR 666/166D as a parent. These 3 families had exceptional bunch indices at the Pamol site :- Family Bunch Index (BI) PK % (107% of trial mean) PK % (116% of trial mean) PK % (106% of trial mean) A fourth family, PK116, also had UR666/166D as a parent but was not originally selected as its EOE yield (5.41 t/ha) was 4% below the mean of trial 74/2. It was also tall, the height coming from UR 609/316D, its other parent. It was later added into the selection list due to the presence of UR 666/166D in its pedigree, as well as its good BI (50.0%). PK116 is the only family in the final selection list that is of pure UR origin. the rights to all information contained herein. Page 8 of 26

9 PK119, the self of UR 706/512D, was low yielding, as would be expected for a selfing, but it had good oil/bunch of just below 19% at Pamol, and an exceptional 24.5% at Paya Lang. UR 706/512D was one parent of PK118 and PK106, both the highest yielders (for EOE) in their respective trials. The progeny in the self had a high number of small-ish bunches, consistent with expectations of its Klanang Bharu parents. Rosenquist (1985) noted that this type of palm often transmits high yield. A later visit to Chemara by the senior author and the late Eric Rosenquist in 1989 confirmed the value of UR 706/512D as a source of good seed parents. PK100 was a later addition to the final selection list, originally not considered due to its height and large fronds, despite being ranked 3 rd for EOE yield (5.42 t/ha) in trial 74/1. However, it had an oil/bunch of 19% at Pamol and 22% at Paya Lang, and a high oil/mesocarp. For individual palm selection within the selected families, the following was emphasized at the Pamol site (Donough & Law, 1995) :- high product yields (EOE); highly heritable (thick mesocarp, thin shell) and desirable (high oil/mesocarp, big fruits) fruit characters; desirable vegetative characteristics (short trunk, small petioles, large leaf area, shorter rachis, and high frond production); and high efficiency of dry matter partitioning (high bunch index (BI) and leaf area ratio (LAR)). Table 5 shows an example of the characteristics of individual palms selected at the Pamol site, compared to the plot, family and trial means (data extracted from Rosenquist, 1985). Donough and Law (1995) gave the mean values for some key parameters from 99 individual palms (representing 11.5% of total Ds in the trials) that were eventually selected from 864 Ds at Pamol that data is also reproduced in Table 5. It is worth noting the values for some key parameters particularly oil/mesocarp (trial mean of 46-47%, despite the cold extraction method used at Pamol) and shell/fruit (trial mean of 26-27%) as these appear much better than usually reported even for Deli materials. Also worth noting is the value of bunch index at 49% for the trial mean, again a higher value than would normally be expected. These may be reflective of the criteria and emphasis applied earlier by Dr Jaap Hardon at OPGL when selecting the D parents to make these crosses. At Dunlop, vegetative measurements had been taken, but unlike at Pamol, growth indices not estimated using the data. Instead, visual appearance of individual palms, in terms of growth vigour and size, and appearance of cut fruit sections, were considered along with FFB yield and bunch analysis data during individual palm selection in the field (S B Chiu, personal communication). In this way, 180 palms (representing 20.8% of total Ds in the trial) were eventually selected from the population of 864 duras at Paya Lang. This difference in data consideration and selection emphasis was the beginning of a divergence in the new generation of D palms bred from the original LBP families. It was thus fortunate, from the breeder s point of view, that the LBP D materials at the Pamol site had now been selected for the 2 nd time along similar lines as the initial D parents had been selected by Dr. Hardon 15 years earlier. Unfortunately, it was not possible to compare the Pamol selections against the parent generation due to lack of data from OPGL. the rights to all information contained herein. Page 9 of 26

10 Table 5. An example of individual palm selection of dura seed parents at Pamol 1 (Key characters selected for at individual palm level in bold type) Parameters Considered Trial Mean Family Mean Plot Mean Selected Palms (3) 9 Mean of 99 palms 10 Yield & its components 2 FFB Yield 3 (t ha -1 ) Bunch no. (yr -1 ) Av. Bunch Wt. (kg) Oil/bunch (%) Oil Yield 4 (t ha -1 ) Kernel/bunch (%) EOE Yield 5 (t ha -1 ) Bunch & fruit components 6 Fruit/bunch (%) Mesocarp/fruit (%) Shell/fruit (%) Oil/mesocarp (%) Av. Fruit Wt. (g) Vegetative measurements 7 Rachis length (m) Leaf area (m 2 ) Petiole x-section x (cm 2 ) New fronds (yr -1 ) Palm height (m) Growth indices 8 BI (%) LAR (m 2 kg -1 ) data from trial 74/1, family PK108, plot 22; 2 yield in t/ha extrapolated from single palm data at 138 palms/ha; 3 6 years mean ; 4 extrapolated using oil/bunch & FFB yield; 5 estimated oil equivalent = oil yield + 60% kernel yield; 6 min. 150 analyses/family, cold extraction (Blaak, 1970); 7 data for 1984 (10 years after planting); 8 following methods of Corley & Breure (1981); 9 3 palms selected in the plot of 16; 10 from Table 3 in Donough & Law (1995). Yong and Leong (1998, 2000) referred to the selected individual D palms at Paya Lang as IOI Deli Duras, or IDD. With Pamol now in the IOI Group, the term IDD is used in a collective sense for the rest of this review, i.e. including all individual D parents selected from the LBP trials at both sites. Where there is a need for differentiation, IDD from Pamol will be termed IDDPAM. Since these IDDs were derived from pedigrees proven to give good DxP progeny in the past, the respective stations i.e. Pamol and IOI, used them immediately for seed production. The first controlled pollination seed bunches were harvested at both sites in early 1986, and the first commercial field plantings were established from Performance of the DxP derived from the IDDs will be considered in a later section further below. the rights to all information contained herein. Page 10 of 26

11 First cycle selection from the LBP in IOI Group (b) Pisifera source One of the 2 IBY crosses called D2 at Dunlop and PK111 at Pamol was selected as the better family at both sites, based on the FFB yield, bunch analysis characteristics of the T sibs in it. At that time, the full ancestry of these crosses (see Appendix 2) was not known. It was thus fortunate that the better cross was not the one with Deli genes in its ancestry. Some data for both IBY families from the Pamol site are given in Table 6 below. Both T and D sibs in PK111 (=D2) were shorter, and had a higher yield from a higher number of smaller bunches. Table 6. Performance of two IBY families at Pamol Parameters PK111 PK111 PK111 PK112 PK 112 PK 112 Considered T Mean 1 D Mean 2 P Mean 3 T Mean D Mean P Mean Yield & its components 4 FFB Yield 5 (t ha -1 ) Bunch no. (yr -1 ) Av. Bunch Wt. (kg) Oil Yield 6 (t ha -1 ) EOE Yield 7 (t ha -1 ) Bunch & fruit components 8 Oil/bunch (%) Kernel/bunch (%) Shell/fruit (%) Vegetative measurements 9 Leaf area (m 2 ) ,6 Petiole x-section (cm 2 ) New fronds (yr -1 ) Palm height (m) Growth indices 10 BI (%) LAR (m 2 kg -1 ) mean values for tenera sibs only; 2 mean values for dura sibs only; 3 mean values for pisifera sibs only; 4 6 years mean ( ); 5 yield in t/ha extrapolated from single palm data at 138 palms/ha; 6- extrapolated using oil/bunch & FFB yield; 7 estimated oil equivalent = oil yield + 60% kernel yield; 8 bunch analysis done for tenera sibs only, cold extraction (Blaak, 1970); 9 data for 1984 (10 years after planting); 10 following methods of Corley & Breure (1981). Most of the P sibs in these IBY families were only partially fertile or infertile, so could not be chosen based on their own yields. At Pamol, the 15 Ps in PK111 were ranked on LAR (Rosenquist, 1985). Pollen collection efforts were thus guided by these rankings, eventually resulting in pollen being obtained from 6 Ps ranked 1-3 and 5-7. Data for these 6 palms are given in Table 7. As can be seen in the data presented, the Ps can be classified into 2 groups based on their vegetative dry matter (VDM) production :- Low VDM group palms 21/6, 21/14 and 26/2 High VDM group palms 16/1, 21/8 and 26/7 the rights to all information contained herein. Page 11 of 26

12 Table 7. Vegetative characteristics and growth indices of selected IBY 1 pisiferas at Pamol (Key differences between groups in bold type) Parameters Low VDM 2 Palms High VDM Palms Considered 21/6 21/14 26/2 Means 16/1 21/8 26/7 Means Leaf area (m 2 ) Petiole x-section (cm 2 ) New fronds (yr - 1 ) Palm height (m) VDM (t ha - 1 ) LAR (m 2 kg -1 ) LAR ranking IOI Binga-Yangambi; 2 vegetative dry matter. The difference between the groups arose from the frond production rate, with the low VDM group producing a third less new fronds a year, leading to trunks about a sixth shorter and VDM almost a third lower. At Paya Lang, no use was made of vegetative data to rank the Ps in family D2 (=PK111). Pollen was eventually successfully collected from 7 Ps 73/4, 73/6, 73/14, 75/5, 76/7, 77/10 and 80/9. Although the selected IBY Ps were of known pedigree, they were unproven as parents for DxP seed production, unlike, say, the AVROS (Av) line descended from the pisifera BM119 at Golden Hope s Banting Oil Palm Research Station (OPRS) (see Appendix 3). The selected IBY Ps were thus used to produce DxP test crosses with IDD parents for evaluation in field trials prior to their use as P parents for commercial DxP seed production. At Pamol, palm 21/6 was first to be tested in 1987 (trial DP5), but the main testing was carried out only later in 1991 (trial DP10 palms 16/1 and 21/8) and 1992 (trial CBP13 all except palm 21/6). Some of the results from CBP13 have been reported recently (Lim et al, 2003; Teo et al, 2004) and will be reviewed in a later section further below. The Paya Lang P selections were tested in 1988 (trial O/10-5/PL palms 73/4, 73/14, 75/5, 76/7 and 80/9) and 1990 (trial O/10-10/PL palms 73/6, 77/10 and 80/9). Results were reported by Yong & Leong (1998, 2000) and will be reviewed further below. the rights to all information contained herein. Page 12 of 26

13 First cycle seed production of the LBP selections in IOI Group For ease of discussion, the production centres or sites shall from here on be called IOI RC and Pamol. The 1 st cycle IDD parents were used until 1996, after which they were replaced by new selections from the 2 nd cycle. The last commercial plantings of 1 st cycle seed were in From the previous section, we showed how the LBP-derived selections in the two centres had diverged at the point of selection of the IDD parents. At the start of seed production, the respective programmes diverged further in respect of the pollen parents used. In the following discussion, we will consider seed production from each site separately. 1. Seed production from the Paya Lang selections by IOI RC The first DxP bunch from controlled pollination between IDD and IBY parents was harvested in April When Dunlop became part of the IOI Group in 1990, this first DxP seed was named IOI 101. Collaboration with AAR a marriage of convenience From very early on, it was recognized that while the LBP had produced D parents with good potential, it lacked a source of P parents with proven pedigree for DxP seed production. As luck would have it, AAR at that time had proven Ps of Dumpy-AVROS (Dy-Av) origins (Soh et al, 1981), but lacked good Ds. A marriage of convenience was thus arranged that allowed IOI to use pollen from AAR s proven Dy-Av Ps for commercial seed production (Yong & Leong, 2000) this seed was called IOI 201 (Chee, 1993). To produce this IOI 201 seed, the LBP trial at Paya Lang was re-visited and the already selected IDD parents were re-examined in 1987, with the aid of Dr A C Soh from AAR, and re-selected (S B Chiu, personal communication). These re-selected IDD parents were also used with the IBY P parents from Paya Lang to produce IOI 301 seed (Chee, 1993; S B Chiu, personal communication), whereupon the production of IOI 101 seed was supplanted. AVROS materials from PORIM In 1991, IOI obtained pure Av material from the Palm Oil Research Institute of Malaysia (PORIM, now incorporated into the Malaysian Palm Oil Board, MPOB) (Yong & Leong, 2000) this material was established in a genetic block (Trial O/10-15/G) at Gomali Estate, Johore. The ancestry of this material traces back to the original selections in Sungai Pancur, Indonesia (lineage given in Appendix 3). They are the 3 rd cycle derivatives from BM119. This material has since been further bred in IOI and the 4 th cycle derivatives have been planted in Sabah (trials O/10-1S/LP2 in 1996 and trial OP/10-4S/LP1 in 1999) and Peninsular Malaysia (trial O/10-41/R in 1999). Pure Av pollen of progeny-tested Ps was also obtained from PORIM at that time for use to produce DxP seeds using the IDDs (Yong & Leong, 2000). This seed was called IOI 401, and replaced the IOI 201. IOI 401 seed was produced until pollen was available in 1995 from selected Av Ps in trial O/10-15/G (the Gomali Av Ps), soon after which its production was discontinued. the rights to all information contained herein. Page 13 of 26

14 Thus, in the 10-year period from the time seed production started in 1985 until Av P pollen become available from Gomali in 1995, IOI RC produced three different types of DxP seed by using different pollen parents : Ybi DxP seed (IDD x IBY origin i.e. pure LBP-derived seed) the IOI 101 seed, supplanted by the IOI 301 in 1987, both produced using pollen from the IBY Ps, initially for use within IOI, later for sale after progeny test results were obtained for the IBY Ps (Yong & Leong, 1998, 2000). This seed was produced throughout the entire period under consideration; Dy-Av DxP seed the IOI 201 seed, using pollen from AAR s proven Dy-Av Ps; this seed was produced between 1987 and Commercial plantings of IOI 201 in IOI s estates started from 1989 (Chee,1993) and continued until 1993, a year or so after the collaboration with AAR ended; and Av DxP seed the IOI 401 seed, using pollen from proven Av Ps from MPOB, from 1991 until After 1995, pollen from the Gomali Av Ps was used to produced IOI 501. This 3 rd type of seed IOI 401 and IOI 501 is of the same genetic stocks as that produced at Pamol (see below). 2. Seed production from the Pamol selections Eric Rosenquist, at the time a consultant to Unilever Plantations Group and Harrisons & Crosfield for their Combined Breeding Programme (CBP), helped broker a deal that allowed Pamol to purchase pollen of proven Av Ps from the Dami OPRS in Papua New Guinea. The performance of DxP seed produced using the Dami Av Ps in Malaysia has been reported by Donough and Law (1995). This arrangement was to last until 1996, following which Av pollen was obtained from MPOB until present day. Thus at Kluang, Pamol had produced a single type of seed IDDPAM x Av, or Pamol seed throughout the period under consideration. This Pamol seed was of the same genetic origins as IOI 401 and IOI 501. The differences were :- The selection method for IDDPAM, as described earlier; and The generation of Av P the Dami Av Ps were the 1 st cycle after BM119, whereas the MPOB Av used to produce IOI 401 were the 2 nd cycle after BM119. The Gomali Av Ps used for IOI 501 production were the 3 rd cycle after BM119 (see Appendix 3). the rights to all information contained herein. Page 14 of 26

15 Performance of materials derived from the first cycle LBP selections in IOI Group Seed was produced using IDDs from the 1 st cycle LBP selections between 1985 and The bulk of commercial scale planting started in 1988 and continued to 1997, with 1998 a transition year between the 1 st and 2 nd cycle materials. Using data within the period , we will now consider the performance of the 1 st cycle LBP derivatives from the the following aspects :- How good were the IBY Ps? How did pure LBP-derived seed, i.e. IOI 301, compare with seed of other genetic origins including IDDPAM x Av seed, i.e. Pamol seed? How did Pamol seed compare to other Deli x Av seed? 1. Progeny test results for IBY Ps Yong & Leong (1998) published the first results from a progeny test trial O/10-10/PL planted in 1990 in which 3 IBY Ps selected from Paya Lang palms 73/6, 77/10 and 80/9 were tested. Results from an earlier trial, O/10-5/PL, planted in 1988 in which 5 IBY P selections from Paya Lang palms 73/4, 73/14, 75/5, 76/7 and 80/9 were tested were given in a later report (Yong & Leong, 2000). In the later paper, the results for only 4 of the 5 IBY Ps tested were included results for one of the Ps, palm 75/5, were excluded as it was represented by just a single cross. In summary, we can conclude the following from the results reported by Yong and Leong (1998, 2000) :- All 6 IBY Ps produced DxP progeny that easily surpassed the specifications for seed parents as defined in Malaysian Standard MS157:1997, and IBY palm 80/9 gave DxP progenies with superior oil/bunch and oil yields some 10% better than the other 4 Ps; the next best IBY P was palm 76/7 which produced DxP progeny with the highest oil/bunch from fruits with thicker mesocarp and thinner shell. A control cross from MPOB was used in trial O/10-10/PL. This cross (0.175/964D x 0.151/2626P) was between an Elmina Deli dura and a pisifera from the MPOB Nigerian germplasm collection, and routinely supplied by MPOB to oil palm breeders for use as a link cross between trials. Unfortunately, in both papers by Yong and Leong, this control cross was wrongly declared a Deli x Av cross. The comparisons made by Yong and Leong between IOI 301 and the wrongly supposed Deli x Av control cross were therefore invalid. That aside, the results directly related to the performance of the IBY Ps as seed parents in their own right were correct and valid. At Pamol, 5 of the 6 IBY Ps were tested in a large progeny test programme involving a comparison of Ps of various origins, crossed with Ds having varying levels of Deli genes. This programme, called CBP13, was conceived by Rosenquist (1988). Initial results from the programme were recently published (Lim et al, 2003; Teo et al, 2004). Data extracted from these two latest papers, and from two earlier trials where the 6 th IBY P, and 2 of the others, were tested, are summarized in Table 8 for a comparison of the IBY Ps selected from the Pamol site. the rights to all information contained herein. Page 15 of 26

16 Table 8. Comparative performance of 6 IBY 1 Ps from Pamol against Av P 2, expressed by their DxP progeny in 5 field trials 3 Parameters Low VDM 4 palms High VDM palms IBY Av Considered 21/6 5 21/14 26/2* 16/1 21/8* 26/7* Mean Mean 6 Yields 7 (all IBY data as % of Av; all Av data as % of trial mean) FFB yield (kg palm -1 ) Oil yield (kg palm -1 ) EOE yield (kg palm -1 ) Bunch & fruit characteristics (All actual values shown) Fruit/bunch (%) Mesocarp/fruit (%) Shell/fruit (%) Oil/mesocarp (%) Oil/bunch (%) Kernel/bunch (%) Av Fruit Wt. (%) Vegetative measurements 8 (All IBY data as % of Av; all Av data as % of trial mean) Rachis length (m) Petiole x-section (cm 2 ) Leaf area (m 2 ) New fronds (yr -1 ) Height incr (cm yr -1 ) Trunk width (cm) Growth indices 9 (All IBY data as % of Av; all Av data as % of trial mean) Bunch Index (%) VDM (kg palm -1 ) Leaf area ratio (m 2 kg -1 ) P LAR rank IBY=IOI Binga-Yangambi; 2 Av=AVROS; 3 data from 5 field trials, trial 1 planted 1987 (Donough & Law, 1995 detailes in Table 6), trial 2 planted 1991, trials 3-5 planted (details in Lim et al, 2003); 4 vegetative dry matter (see Table 7 above for details of palm ranks); 5 palm 21/6 tested in trial 1, palm 16/1 in trial 2 & 3, palm 21/8 in trials 2, 4 & 5, other in trials 4-5; 6 Av means all as % of trial mean for yields and vegetative measurements, actual data for bunch analysis; 7 data for trial 1 Nov. 92-Oct. 96, trial 2 Jun. 94-May 01,others from trials 2-4 Jul. 96-Jun. 00; 8 data for trial 1 taken Nov. 94 (7 th year), others from trials 2-5 at 7½ years old; 9 following methods of Corley & Breure (1981); 10- original ranking assigned to each P based on leaf area ratio (LAR) see table 7 above. * Palms 21/8, 26/2 & 26/7 highlighted as best of the IBY Ps. In summary, the IBY Ps generally produced DxP progeny as good as Av Ps for FFB yield, but lost out on oil and EOE yields due to poorer fruit characteristics and oil/mesocarp. Progeny of the IBY Ps were generally 10% shorter than Av P progeny (except those derived from palm 26/2) but somewhat fatter, with more massive and longer fronds. the rights to all information contained herein. Page 16 of 26

17 However, there was considerable variability between the IBY Ps. Palm 26/2, and palms 21/8 and 26/7, produced DxP progeny that outyielded Av P progeny for EOE by 30% and 10%, respectively. In the case of palm 26/2, this was achieved through better competitiveness its progeny were massive in size, even height, compared to Av P progeny. Palms 21/8 and 26/7, on the other hand, produced DxP progeny that were 10% shorter than Av P progeny. Even palm 16/1 produced DxP progeny as good as Av P for EOE yield, but 15% shorter. For a wider analysis, the reader should refer to the papers by Lim et al (2003) and Teo et al (2004), who compared the IBY Ps against Ps from other origins, including Ekona and URT. PK111 (=D2) is being further developed in the IOI breeding programme. The 6 IBY Ps selected from Pamol were earlier classified into 2 sets on the basis of their VDM (see Table 7 earlier), but there appeared no reflection of this in their DxP progeny. Similarly, the earlier ranking of the IBY Ps by LAR also did not appear to be reflected in the results obtained in their DxP progeny. There was no difference in rate of new frond production in the DxP progeny as had been earlier observed for the IBY Ps. However, if palm 26/2 was excluded, then it appears that the high VDM Ps did produce DxP progeny with a higher VDM, that were somewhat taller with thicker trunks. 2. IOI 301 and Pamol seed vs other Malaysian seed Yong and Leong (2000) presented results from a 1988 DxP source comparison trial, O/10-6/PL, showing that IOI 301 compared favourably with 4 other Malaysian seed based on records up to Pamol seed was not included in this trial. More data are available now, and Table 9 shows results up to June Table 9. Results 1 from trial O/10-6/PL 2 comparing IOI 301 against 4 other Malaysian seed FFB Oil/ Oil Kernel EOE Seed Yield 3 Bunch 4 Yield 5 /bunch Yield 6 IOI Source A Source B Source C Source D Trial Mean All data from IOI RC; 2 Trial established 1988, Paya Lang Estate, at 136 palms ha -1, Batu Anam/Durian Series; 3 yield in kg palm -1 annualized for period March 1991-June 2003; 4 in %, bunch analysis as described by Rao et al (1983); 5 in kg/palm, product of annual FFB yield and oil/bunch; 6 in kg palm -1, estimated oil equivalent (oil yield + 60% kernel yield). the rights to all information contained herein. Page 17 of 26

18 On the basis of the data presented, there was little difference between IOI 301 and the other seeds. For FFB yield and oil/bunch, there was a mere 5-6% difference between the highest (Source B) and lowest (Source C) ranks, with IOI 301 ranked 2 nd and joint third for both parameters respectively. The difference between the highest and the lowest for oil and EOE yield was about 10-11%, with IOI 301 ranked 2 nd, only 3% below Source B, in both cases. In this instance, Source B seed was of Deli x Av origins, thus the results agreed with the earlier P comparisons. Source C seed was of Deli x Ybi origins, similar to IOI 301, but ranked lower for all parameters except kernel/bunch. Lim et al (2003) and Teo et al (2004) had also found that the IBY Ps compared better against other P of Binga (i.e. Ybi) origins. IOI 301 was included in another trial comparing various Malaysian DxP seed that was conducted in Sandakan by Jabatan Pertanian Sabah (JPS). This trial was established in late-1989 at the JPS Agricultural Research Station at Ulu Dusun (UDARS) and included seed from 9 Malaysian sources, classified by their P origins as follows :- Deli x Av P 3 sources (Pamol and 2 others) Deli x Ybi 3 sources (IOI 301 and 2 others) Deli x Dy Av 1 source Deli x UR or NIFOR (Nf) 2 sources Data for FFB yield, bunch analysis and vegetative measurements from this trial were kindly provided by Betty Kwan (oil palm agronomist at UDARS) and a summary is presented in Table 10. We calculated growth indices from the data provided and key parameters are also shown in Table10. In this trial, it was clear that DxP material derived from the IDDPAM selections were better than all other seed, including IOI 301, for yield of oil and EOE. The fruit characteristics of the IDDPAM x Av (i.e. Pamol) seed were even better than the other 2 Deli x Av seed sources, suggesting that the strong emphasis given to oil yield characteristics during selection of IDDPAM had some influence over and above that of Av P. Similarly, the vegetative characteristics of the Pamol seed also differed from the other Av seeds, suggesting that the IDDPAM may have had an influence. Bunch index in the Pamol seed was better than the other 2 Av seeds, VDM was lower and LAR far higher. Given that the Pamol seed differed from the other 2 Av seeds in the way described above, the difference in vegetative characteristics between the Pamol seed and IOI 301 were likely to be more reflective of the different bias in criteria emphasis during the selection of IDDs at Paya Lang, although the male parent contribution also could not be entirely ruled out. Selection in the IDDPAM for smaller petioles and higher rate of new frond production appeared to be reflected in the Pamol seed. A feature of both the Pamol seed and IDD 301 that also stood out in this trial was the large fruit size. The Pamol seed gave more oil and EOE yield than the other seeds at similar levels of FFB yield, clearly indicating that it was more profitable for the grower. Isa et al (2005) are presenting results in this Seminar from the 3 rd round of Malaysian DxP seed comparison coordinated by MPOB. This series of trials were established in 1996 and both IOI 301 and Pamol seed were represented, along with 7 other seed sources, 5 of which were commercial seed producers. These trials were the last in which materials derived from the 1 st cycle LBP selections were represented, as after 1997, 2 nd cycle selections had taken over. the rights to all information contained herein. Page 18 of 26

19 Table 10. Results from a trial 1 comparing 9 Malaysian seeds in Sandakan, Sabah Parameters Considered PAM Other IOI Other Other Dy Trial Av 2 Av Ybi 5 UR/Nf 6 Av 7 Mean 8 Yield 9 (All data as % of trial mean) FFB yield (t ha -1 ) Oil yield (t ha -1 ) EOE yield (t ha -1 ) Bunch & fruit characteristics 10 (actual values shown) Fruit/bunch (%) Mesocarp/fruit (%) Shell/fruit (%) Oil/dry mesocarp (%) Oil/bunch (%) 29, Kernel/bunch (%) Av Fruit Wt. (%) Vegetative measurements 12 (All data as % of trial mean) Rachis length (m) Petiole x-section (cm 2 ) Leaf area (m 2 ) New fronds (yr -1 ) Height incr (cm yr -1 ) Trunk width (cm) Growth indices 13 (calculated from above data provided, shown as % of trial mean) BI (%) VDM (t ha -1 ) LAR (m 2 kg -1 ) trial at Jabatan Pertanian Sabah s Ulu Dusun Agricultural Research Station, Sandakan, planted 1989; 2 IDD x Dami Av seed from Pamol; 3 2 other Deli x Av seed sources; 4 IDD x IBY seed; 5 2 other Deli x Ybi seed sources; 6 2 seed sources using UR &/or Nifor Ps; 7-1 seed source; 8 all data shown for trial mean are actual values; 9 FFB yield from ; 10- bunch analysis from , teneras only; 11 other tables show oil/wet mesocarp; 12 measurements done in 2001 (12 th year after planting); 13 calculated from data provided using methods of Corley & Breure (1981). The results of Isa et al (ibid) add further credence to those we had presented in the foregoing the Pamol seed, of IDDPAM x Av origins, gave high oil yields consistently across sites, with its superiority arising from its excellent fruit characteristics giving it the highest oil/bunch exceeding 29%. If there was a drawback in this series of trials, it was that 2 of the largest seed producers in Malaysia Chemara (including Highlands Research Unit, HRU) and Golden Hope did not participate. However, their absence does not invalidate the results obtained. the rights to all information contained herein. Page 19 of 26