GeneticPenelitian Green Pertanian Super RiceTanaman Resistance Pangan .... (Susanto et al.) Vol. 3 No. 3 Desember 2019: 111-116 DOI: http//dx.doi.org/10.21082/jpptp.v3n3.2019.p111-116

Genetic Variability and Heritability of Green Super Rice Resistance to Rice Ragged Stunt Virus and Rice Grassy Stunt Virus

Variabilitas Genetik dan Heritabilitas Ketahanan Galur-galur Green Super Rice (GSR) terhadap Virus Kerdil Hampa dan Virus Kerdil Rumput

Untung Susanto1*, Nafisah1, Wage R Rohaeni1, Baehaki SE1, Sarlan Abdulrahman1, dan Jauhar Ali2 1Indonesian Center for Rice Research (ICRR) Street 9 Sukamandi, Subang,West Java, 41256 *E-mail: [email protected] 2International Rice Research Institute (IRRI) Los Banos, Laguna,

Naskah diterima 08 Agustus 2019, direvisi 28 November 2019, disetujui diterbitkan 2 Desember 2019

ABSTRAK ABSTRACT

Variabilitas genetik dan heritabilitas galur-galur Green Super Rice Brown planthopper (BPH) is the main pest of rice in Indonesia, (GSR) tahan terhadap virus kerdil hampa dan virus kerdil rumput. occurs every year with the tendency of increasing. BPH serves as Wereng batang coklat (WBC) merupakan salah satu hama utama a vector of Rice Ragged Stunt Virus (RRSV) and Rice Grassy Stunt tanaman padi di Indonesia. Serangan WBC selalu terjadi setiap Virus (RGSV). Green Super Rice (GSR) is designed to resist major tahun dengan luasan yang berfluktuasi. Selain sebagai hama, pests and diseases, so that it needs less pesticide to save the WBC juga merupakan vektor virus yang menyebabkan penyakit quality of environment. GSR was developed at IRRI and in kerdil rumput (Rice Grassy Stunt Virus, RGSV) dan kerdil hampa and had been tested in Indonesia from 2009 until 2019. This research (Rice Ragged Stunt Virus, RRSV). Green Super Rice (GSR) dirakit was aimed to study the genetic variability and heritability of 26 untuk tahan terhadap hama dan penyakit utama tanaman padi, selected GSR lines resistant to RRSV and RGSV. The experiments agar aplikasi pestisida lebih sedikit untuk lebih ramah terhadap were conducted at Experimental Station of ICRR Sukamandi and lingkungan. GSR pertama dirakit di IRRI dan China dan galur- Pusakanagara. The treatments were arranged in a randomized galur yang dihasilkan diuji di Indonesia sejak tahun 2009. Penelitian complete block design with three replications. Transplanting was ini bertujuan untuk mempelajari variabilitas genetik dan done using 21 days old seedling at 25 cm x 25 cm plant spacing on heritibilitas sifat tahan terhadap RRSV dan RGSV 26 galur GSR 1 m x 1 m plot. BPH and virus infestation occurred naturally due to terseleksi, bersama dengan empat varietas cek. Percobaan the BPH outbreak during the Dry Season (DS) of 2010. Percentage dilaksanakan di Kebun Percobaan BB Padi di Sukamandi dan of plants showing RRSV and RGSV symptom was measured as Pusakanagara, menggunakan rancangan acak kelompok tiga criteria for the resistance of the genotype to the viruses. Results ulangan. Tanam pindah dilakukan menggunakan benih umur 21 showed that the tested genotypes indicated high genetic variability hari, pada plot berukuran 1 x 1 m2 dengan jarak tanam 25 x 25 cm. on the virus’s resistance and the heritability was classified as Serangan WBC disertai infestasi virus terjadi secara alami, karena medium for the RGSV resistance. The genotypes showed low epidemi hama WBC terjadi sepanjang musim (Musim Kering 2010). genetic variability and low heritability on the resistance to RRSV. Persentase tanaman yang menunjukkan gejala serangan RRSV This information implied that breeder is possible to develop resistant dan RGSV digunakan sebagai kriteria tingkat ketahanan genotipe lines to RGSV. Genotype HUANGHUAZHAN and HHZ 12-Y4-Y3- terhadap kedua virus. Hasil penelitian menunjukkan bahwa Y1 are consistently resistant to RGSV so that those genotypes genotipe yang diuji memiliki keragaman yang tinggi dan could be used as donor gene for further plant breeding activities. heritabilitas medium untuk sifat tahan terhadap RGSV, tetapi Selection of resistant individual plants in the breeding population is menunjukkan keragaman genetik dan heritabilitas rendah untuk recommended to be done at earlier generations for RGSV and at sifat tahan terhadap RRSV. Hal ini mengindikasikan adanya peluang more advanced generation for RSSV. untuk merakit varietas tahan terhadap RGSV. Galur Keywords: Green Super Rice, genetic variability, heritability, HUANGHUAZHAN dan HHZ 12-Y4-Y3-Y1 secara konsisten tahan RRSV, RGSV. terhadap RGSV sehingga dapat digunakan sebagai donor gen dalam kegiatan pemuliaan tanaman lebih lanjut. Seleksi ketahanan dari populasi persilangan disarankan dilakukan pada generasi awal untuk sifat tahan RGSV dan generasi lanjut untuk sifat tahan RSSV. INTRODUCTION Kata kunci: Green Super Rice, keragaman genetik, heritabilitas, kerdil rumput, kerdil hampa. Brown Plant Hopper (BPH, Nilaparvata lugens) is a major rice pest in Indonesia. BPH has regularly caused great

111 Penelitian Pertanian Tanaman Pangan Vol. 3 No. 3 Desember 2019: 111-116 losses in rice production in Indonesia, southeastern and genotype to RGSV without having penalty on any far eastern Asia, including China, , Thailand, agronomical performance. , Pakistan, Malaysia, Philippines, , Indonesia, Ta et al. (2013) showed that there was no relation and also Korea. BPH attack in Indonesia had increased between genetic diversity and the geographic 5-folds from 47,473 ha in 2009 to 218,060 ha in 2011, but distribution of RGSV isolates. On the other hand, the infested area fluctuated every year (Baehaki 2012; Suprihanto et al. (2015) reported that the nucleotide Baehaki and Mejaya 2014). sequence of RGSV sampled from Java had a strong Aside from being a serious pest by itself, BPH is also affinity with AF486811-Philippines. Dini et al. (2015) also acting as a vector for RRSV (Rice Ragged Stunt Virus) reported that RRSV isolate of Subang had high homology and RGSV (Rice Grassy Stunt Virus). BPH outbreak is (97.1%) with RRSV isolates of Vietnam, Philippines, and frequently followed by virus attack. RGSV was first Thailand, while RGSV isolates of Subang had high reported in Indonesia in 1970 and RRSV in 1997 (Du et homology (95.8%) with RGSV isolate of Longan, Vietnam. al. 2005). RGSV attack on rice induces leaf yellowing, Green Super Rice was developed to have a high yield stunting, and excess tillering (Satoh et al. 2013). Nguyen and stable under optimum and low input conditions, as et al. (2015) reported that there were two main viruses well as it had some degree of resistance to major pests causing the disease in rice: the Rice Grassy Stunt Virus and diseases (Zhang 2007). It purposedly contained a (RGSV) and Rice Ragged Stunt Virus (RRSV). Dini et al. wide genetic background of resistance genes to major (2015) further confirmed that based on specific DNA pests and diseases. Among the GSR materials, it is sequence of the coating protein from the virus taken expected to be found some lines that are resistant to from Sukamandi in November to December 2014, the RGSV and RRSV. The GSR materials were selected for RGSV and RRSV were identified. high grain yield and acceptable amylose content for Outbreak of these two viruses had decreased rice Indonesian rice consumers (Susanto and Jumali 2011). production in Indonesia and in Vietnam. In 2010, the Iswanto (2015) reported that heritability of resistance to BPH attack which was followed by RRSV and RGSV RGSV and RRSV was medium where selection could be infection caused a decline in rice production of around conducted at the early generation. IR70213-10 IR-CPA 4- 1 million tons (Baehaki 2012). Suprihanto et al. (2015) 2-3-2, IR-RRRR-75 417 457-2, PSB RC 68, IR 82810-407, reported that the diseases that attacked rice crop in the Swarna, Mahsuri, and IR70213-10 IR-CPA 2-3-2-1 were planthopper infested regions were two kinds, i.e. Rice identified as tolerant to RGSV and RRSV under an ragged stunt virus (RRSV) and Rice grassy stunt virus epiphytotic conditions of these diseases. Therefore, the (RGSV). RGSV and RRSV mostly occurred after the BPH resistant lines would be expected to be readily adopted was controlled by spraying a pesticides. RGSV in by farmers upon release. As it was intended, GSR lines Indonesia is considered as type 2, found in Java and in are also expected to be well adapt to various North Sumatra (Baehaki and Mejaya 2014). RRSV dan agroecosystems. RRSV viruses in Indonesia, especially RGSV control by applying pesticides was reported in in Java showed similarities with viruses found in other effective (Baehaki et al. 2017). The most economical and parts of the world. Kusuma et al. (2018) reported that by effective way of controlling RRSV diseases, if available is using specific primer for coat protein (CP) gene, RRSV the use of resistant rice varieties (Wang et al. 2017). This from Indonesia (Yogyakarta, Central Java, and Bali) had technology is considered more cost-effective, 99% similarity with RRSV isolates from Chanting, China. sustainable, and can reach remote areas. Comparing the GSR lines to various check varieties BPH resistant variety could give an escape to rice was expected to identify the superiority of the lines plants from the viruses, although it may break down compared to the check. One of the check varieties, Situ once the gene for BPH resistance is broken. RSSV and Bagendit, was reported as not preferred by BPH, has RGSV resistant variety reduces yield loss due to the plant’s medium resistance to RGSV and has low transmission capability to withstand virus infection, even though the ability of the virus (Suprihanto et al. 2016). BPH had attacked the plants (Suprihanto et al. 2016). The objectives of the research were to evaluate the BPH resistance to insecticides had been reported as one genetic variability and heritability of resistance, to RRSV of factors contributing to BPH epidemic where the case and RGSV. The information is useful to identify superior most possibly due to the inappropriate insecticides uses GSR lines possessing resistance to RRSV and RGSV. The (Sutrisno 2015). Nuque et al. (1982) reported that based heritability estimate would be useful to ascertain the on the segregation pattern in Oryza nivara population, nature of the genetic factor controlling resistance to BPH RGSV resistance was controlled by a single dominant and RRSV-RGSV. gene. It is, therefore, possible to develop resistant rice

112 Genetic Green Super Rice Resistance .... (Susanto et al.)

MATERIALS AND METHOD Score Symptom The experiment was conducted at two locations, namely Sukamandi and Pusakanagara Experimental Station of 7 11-30% height reduction, 3-4 leaves have the Indonesian Center for Rice Research (ICRR) during ragged/twisted symptom, and some on leaf dry season of 2010. The trial was conducted coincide blade and the leaf sheaths with the outbreak of the two viruses’ diseases and the 9 More than 30% height reduction, most leaves vector. have ragged/twisted leaf symptoms, vein The experiment was arranged in a randomized swelling commonly on leaf sheaths and leaf complete block design (RCBD) with three replications. blades Twenty-six GSR lines were evaluated for their genetic variability and their heritability for resistance to RRSV and Disease infection was observed at the heading stage RGSV. Four rice varieties including those resistant varieties for all plants in each plot. Disease incidence was to RRSV and RGSV, namely: Ciherang, Inpari 10, Inpari calculated using the following formula: 11, and Situ Bagendit were used as checks. Transplanting was done using 21-day old seedling at 25 cm x 25 cm n DI = x 100% plant spacing in 1 m x 1 m plot. Crop maintenance was N optimally carried out. Fertilizer with a dosage of 200 kg Urea, 100 kg SP36 and 100 kg KCl per ha was applied, DI = disease incidence where the nitrogen was split into three application times, n = number of infected plants i.e. 10 days, 21 days after transplanting and at flowering N = number of total plants observed initiation. BPH and virus infestation occurred naturally Data were transformed into arch-sin and the due to the BPH outbreak during the season. Percentage variance components were analyzed to separate of plants showing RRSV and RGSV symptom was used genotypic and phenotypic variance to be furthered as criteria for the resistance plant to the viruses, based calculated genetic variance and heritability, as outlined on the Standard Evaluation System for Rice (IRRI 2013) below. as follows. Source of df Mean Expected Score Symptom variation square mean square Replicates r-1 RGSV (type 2) 2 2 Genotypes g-1 M1 r g +  e 1 No symptom observed Residual (r-1)(g-1) M2 2 3 Pale green and slightly narrow leaves, no e height reduction and with numerous small Notes: tillers df = degree of freedom, r = replication; g = genetic 5 Distinct yellow and narrow leaves, 1-10% height reduction, and with numerous small Environmental variance (2 ) = error variance tillers e 7 Yellow to orange and narrow leaves with Genotypic variance (2 ) = some rusty spots, 11- 30% height reduction, g and with few small tillers 2 2 2 Phenotypic variance ( P) =  g +  e 9 Yellow to orange and narrow leaves with numerous rusty spots, >30% height Deviation of variance: reduction and with few small tillers Genetic variability: it was considered as wide if 2 > 2 RRSV g 1 No Symptom observed and vice versa. 3 0-10% height reduction, no ragged/twisted leaf, small and very few veins swelling usually Heritability (h ) = (Allard 1999). The heritability was on leaf collar bs 5 0-10% height reduction, 1-2 leaves have classified as high if it is higher than 50%, medium 20% - ragged/twisted leaf, small and very few vein 50%, and low below 20% (Mc. Whirter 1979). swelling usually on leaf collar

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RESULTS AND DISCUSSION resistance. This trait might be useful for indirect selection of RGSV resistance in rice. Brown planthopper outbreak during 2010 gave a The percentage of RGSV attacked plants in blessing in disguise, of which it was possible to select Sukamandi and Pusakanagara was similar. Genotype resistant variety to RGSV and RRSV pathogens. The Zhonghua 1 had no plant (0%) attacked by RGSV, both in viruses remained in the infected plant even though the Sukamandi and in Pusakanagara. Six other GSR lines host, i.e. brown plant hopper, had been eradicated. with the lowest percentage of RGSV attack from the two During the time, the BPH attack at Sukamandi and locations (Sukamandi and Pusakanagara) were HHZ 11- Pusakanagara was severe, where more than 100 brown Y6-Y1-Y1 (1.33%), HUA 564 (1.33%), HHZ 11-Y11-Y3-DT1 plant hopper insects were found in one plant. Under (2.00%), WEED TOLERANT RICE 1 (2.00%), such a circumstance, it was assumed that all plants were HUANGHUAZHAN (3.33%), and HHZ 12-Y4-Y3-Y1 (4.00%) infected by both RGSV and RRSV viruses. BPH attacked (Table 2). Those seven lines were consistently showed followed by RGSV and RRSV diseases spread to all resistance to RGSV. surrounding rice crop nearby the trials where all plants were finally eradicated, remaining only plants in this trial. These findings also indicated that GSR material Among genotypes, there were variations of surviving contains effective genes controlling resistance to pest plants. The survived healthy plants after BPH chemical and diseases, including bacterial blight (Rohmah et al. control was assumed to represent the resistance 2013), brown spot (Yaqoob et al. 2011), and also weeds genotypes to viruses. Percentage of plant showing a (Chauchan et al. 2015). Some GSR materials were symptom of RGSV and RRSV was considered as reported as drought-resistant (Guan et al. 2010). susceptible to the virus. Resistance to RRSV

Resistance to RGSV Broad sense heritability of the RRSV resistance was found Broad sense heritability of the RGSV resistance among as low, with the value of H was: -0.012 (Sukamandi) and the tested genotypes was classified as medium, the value -0.056 (Pusakanagara). Its genetic variability of this trait of heritability estimates H, was 0.497 (Sukamandi) and was also low (Table 1). Therefore, selecting this trait 0.498 (Pusakanagara) (Table 1). among the tested genotypes might not be feasible. There are some Indonesian local rice varieties had been Genetic variability of the RGSV resistance among the identified as having resistance to RRSV, namely Mentik genotypes was considered wide both in Sukamandi and Wangi, Tetep and Utri Merah (Suprihanto et al. 2016). in Pusakanagara (Table 1), indicating that the tested genotypes had various levels of resistance to RGSV. The genetic variability of the RRSV resistance Resistance genotypes were prospected to be stable genotypes was considered narrow. The occurrence of among regions and overtimes. High heritability of RRSV infection ranged from 0 to 23.33% (Table 2). All resistance suggested that the variation derived from the genotypes had a relatively low incidence of RRSV, genetic factor. Its high variability indicated the possibility indicating that the genotypes relatively tolerant to RRSV. of finding resistant genotypes. Selecting genotype The heritability estimate was also low, suggesting that resistance to virus is tedious work. Indirect selection Selection of resistance to RRSV is better to be conducted based on traits correlated to RGSV resistance would be at advanced generation. Crossing among resistance more practical. Le et al. (2012) reported that oxalate genotypes would be expected to increase the gene content in rice is positively correlated with RGSV frequencies for resistance.

Table 1. Genetic variability and heritability of resistance to RRSV and to RGSV of 26 GSR inbred lines.

2 2 2 2 2  e  g  P hbs Crit. of hbs ( g) 2 x ( g) Gen. variability

Sukamandi RGSV 0.220 0.217 0.437 0.497 Med 0.079 0.158 Wide RRSV 0.056 -0.001 0.056 -0.012 Low 0.009 0.017 Narrow

Pusakanagara RGSV 0.272 0.269 0.541 0.498 Med 0.098 0.196 Wide RRSV 0.047 -0.003 0.045 -0.056 Low 0.007 0.014 Narrow

2 2 2 Note:  g: genotypic variability,  e: environment variability,  P: phenotypic variability, hbs: heritability broad sense

114 Genetic Green Super Rice Resistance .... (Susanto et al.)

Table 2. Percentage of occurring RRSV and RGSV symptom on 26 GSR lines along with 4 check varieties.

RGSV RRSV Genotype Sukamandi Pusakanagara Average Sukamandi Pusakanagara Average

HUA 564 2.67 0.00 1.33 20.00 0.00 10.00 HUA 565 64.00 98.67 81.33 6.67 0.00 3.33 WANXIAN 763 30.67 6.67 18.67 13.33 33.33 23.33 WANXIAN 77 28.00 53.33 40.67 2.67 0.00 1.33 HUANGHUAZHAN 6.67 0.00 3.33 13.33 26.67 20.00 HEXI 41 66.67 98.67 82.67 33.33 0.00 16.67 YUNJING 23 100.00 66.67 83.33 0.00 0.00 0.00 SACG 4 49.33 33.33 41.33 0.00 0.00 0.00 SACG-7 42.67 93.33 68.00 6.67 0.00 3.33 ZHONGHUA 1 0.00 0.00 0.00 20.00 0.00 10.00 WEED TOLERANT RICE 1 4.00 0.00 2.00 16.00 13.33 14.67 WUYUJING 3 52.00 33.33 42.67 9.33 6.67 8.00 WUYUJING 20 33.33 100.00 66.67 42.67 0.00 21.33 BD007 66.67 66.67 66.67 6.67 13.33 10.00 CAU1 74.67 100.00 87.33 9.33 0.00 4.67 CAU2 100.00 93.33 96.67 0.00 0.00 0.00 YUNDAO 1 100.00 98.67 99.33 1.33 0.00 0.67 LUYIN 46 41.33 26.67 34.00 9.33 0.00 4.67 RC8 34.67 33.33 34.00 10.67 0.00 5.33 6527 100.00 100.00 100.00 1.33 0.00 0.67 JH15-1-1-1 8.00 33.33 20.67 13.33 13.33 13.33 D4098 26.67 33.33 30.00 13.33 0.00 6.67 Ciherang 4.00 20.00 12.00 17.33 13.33 15.33 Inpari 10 9.33 0.00 4.67 13.33 0.00 6.67 Situbagendit 6.67 33.33 20.00 16.00 0.00 8.00 Inpari11 4.00 53.33 28.67 20.00 0.00 10.00 HHZ 5-Y3-SAL3-DT1 9.33 0.00 4.67 16.00 0.00 8.00 HHZ 11-Y6-Y1-Y1 2.67 0.00 1.33 22.67 0.00 11.33 HHZ 11-Y11-Y3-DT1 4.00 0.00 2.00 14.67 0.00 7.33 HHZ 12-Y4-Y3-Y1 8.00 0.00 4.00 14.67 13.33 14.00

Notes: Analysis of variance using arc.sin conversion from the percentage data

An alternative approach to enhance the resistance ACKNOWLEDGEMENT to RRSV was reported by developing transgenic rice capable of producing Pns6, a putative movement protein This research was collaboratively funded by GSR Project of RRSV, which acted as a silencing suppressor (Wu, et supported by Bill and Melinda Gates Foundation and al. 2010; Jia et al. 2012). Nguyen et al. (2015) had the Government of the People Republic of China. Untung proposed RNAi strategy as an alternative method to Susanto conducted the experiment and prepared the develop resistance to RGSV and RRSV. Those two report. approaches however, has not been realized. REFERENCE

CONCLUSIONS Allard RW. 1999. Principles of Plant Breeding, 2nd ed. John Wiley & Sons. New York. 485 p. The 26 GSR tested genotypes showed wide genetic Baehaki, S.E., E.H. Iswanto, D. Munawar, Y.H. Song, I.R. Choi, and variability and medium heritability, on resistance to RGSV. H.H. Park. 2017. Abilities of brown planthopper immigrant However, the genetic variability and heritability on transmits Rice Ragged Stunt Virus on rice of some districts of resistance to RRSV were low. Genotype Zhonghua 1, HUA Java-Indonesia. Sch. J. Agric. Vet. Sci. 4(8):300-310. 564, HHZ 11-Y11-Y3-DT1, WEED TOLERANT RICE 1, Baehaki, S.E., M.J. Mejaya. 2014. Wereng cokelat sebagai hama HUANGHUAZHAN, and HHZ 12-Y4-Y3-Y1 are consistently global bernilai ekonomi tinggi dan strategi pengendaliannya. IPTEK Tanaman Pangan 9(1):1-12. resistant to RGSV and thus could be used as donor genes Baehaki, SE. 2012. Perubahan pengendalian hama terpadu (PHT) for resistance to RGSV. konvensional menuju PHT biointensif. In. Hermanto,

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Sunihardi, H. Radianto (eds.). Pros. Sem. Nas. Inovasi Rohmah, N.A., U. Susanto, and T. S. Kadir. 2013. Resistance of Teknologi Berbasis Ketahanan Pangan Berkelanjutan. Green Super Rice (GSR) Lines to Bacterial Leaf Blight. Puslitbangtan. Buku 2. Bogor. Pp. 203-214. Proceeding International Seminar. Technology Innovation for Chauchan, B.S., J. Opena, and J. Ali. 2015. Response of 10 elite Increasing Rice Production and Conserving Environment ‘Green Super Rice’ genotypes to weed infestation in aerobic Under Global Climate Change. ISBN : 9/8-9/9-540-0/2-1. IRRI rice systems. Plant Prod. Sci. 18(2): 228-223. - Indonesian Center for Rice Research. Indonesia Agency for Agricultural Research and Development Ministry of Dini, A.F.B., I.W. Winasa, and S.H. Hidayat. 2015. Identifikasi virus Agriculture Republic of Indonesia. penyebab penyakit kerdil pada tanaman padi di Sukamandi, Jawa Barat. J. Fitopatol, Indon. 11(6): 205-210. DOI: 10.14692/ Satoh, K., K. Yoneyama, H. Kondoh, T. Shimizu, T. Sasaya, Ry. Choi, jfi.11.6.205. K. Yoneyama, T. Omura, and Sh. Kikuchi. 2013. Relationship between gene responses and symptoms induced by Rice Du, P.V., Cabunagan RC., and Choi I-R. 2005. Rice “yellowing grassy stunt virus. Front. Microbiol. 4: 1-10. syndrome” in the Mekong River delta. Omonrice 13:136-139. Suprihanto, S. Somowiyarjo, S. Hartono dan Y.A. Trisyono. 2016. Guan, Y.S., R. Serraj, S.H. Liu, J.L. Xu, J. Ali, W.S. Wang, E. Venus, Preferensi wereng batang cokelat terhadap varietas padi dan L.H.Zhu, and Z.K. Li. 2010. Simultaneously improving yield ketahanan varietas padi terhadap virus kerdil hampa. under drought stress and non stress condition: a case study Penelitian Pertanian Tanaman Pangan 35(1): 1-8. of rice (Oryza sativa L.). J. of Exp. Bot. 61(15): 4145-4156. th Suprihanto, S. Somowiyarjo, S. Hartono, and Y.A Trisyono. 2015. IRRI. 2013. Standard Evaluation System for Rice. 5 Edition. IRRI, Identification and molecular diversity of rice ragged stunt Philippines. 55 pp. virus and rice grassy stunt virus in java, Indonesia. Internat. J. Iswanto, H.E. 2015. Genetic variation and heritability of resistances Sci.: Basic and Applied Research 24(5): 374-386. rice to ragged stunt and grassy stunt virus. AGRIS. http:// Susanto, U. and Jumali. 2011. Karakteristik Mutu Tanak Galur- agris.fao.org/agris-search/search.do?record galur Green Super Rice (GSR) Terseleksi di Indonesia. In T. ID=ID2015010128 Agung, Suwarto, N.D. Sasongko, A.H. Susanto, Tj. Winanto, Jia D, N. Guo, H. Chen, F. Akita, L. Xie, T. Omura, and T. Wei. 2012. dan A. Riyanto (Eds.). Proc. Seminar Nasional Pemuliaan Assembly of the viroplasm by viral non-structural protein Berbasis Potensi dan Kearifan Lokal Menghadapi Tantangan Pns10 is essential for persistent infection of rice ragged stunt Globalisasi. PERIPI Komda Banyumas, LPPM Univesitas virus in its insect vector. Journal of General Virology 93: 2299- Jenderal Soedirman, ISBN 9786021923900. 2309. Sutrisno. 2015. Resistensi wereng batang cokelat padi, Nilaparvata Kusuma, A.F., S. Sulandari, S. Somowiyarjo, and S. Hartono. 2018. lugens stål terhadap insektisida di Indonesia. Jurnal Molecular Diversity of Rice Ragged Stunt Oryza virus in Java AgroBiogen 10(3):115-124. and Bali, Indonesia. B. Life and Enviro. Sci. 55(1): 57-64. Ta, H.A, D.P. Nguyen, S. Causse, T.D. Nguyen, V.V. Ngo, and E. Le, T.T., Luong, V. D., Ngo, T.T.N. and V.K. Pham. 2012. Induced Hébrard. 2013. Molecular diversity of rice grassy stunt virus systemic resistance against rice grassy stunt virus–a in Vietnam. Virus Genes 46(2):383-6. doi: 10.1007/s11262- promising field for ecological rice production. J. Viet. Env. 012-0845-3 2(1): 48-53. Wang, Z.,L. Yu, L. Jin, W. Wang, Q. Zhao, L. Ran, X. Li, Z. Chen, R. Mc. Whiter, K.S. 1979. Breeding of cross-pollinated crops. In R. Guo, Y. Wei, Z. Yang, E. Liu, D. Hu, B. Song. 2017. Evaluation Knight (ed.) plant breeding. Australian Vice Counsellors of rice resistance to southern rice black-streaked dwarf virus Committee. Brisbane. and rice ragged stunt virus through combined field tests, Nguyen TD, Lacombe S, Bangratz M, Ta HA, Vinh do N, Gantet P., quantitative real-time pcr, and proteome analysis. Viruses and Brugidou C. 2015. P2 of Rice grassy stunt virus (RGSV) 9(2): 37. doi: 10.3390/v9020037. and p6 and p9 of Rice ragged stunt virus (RRSV) isolates Wu, J. Du, C. Wang, L. Cai, M. Hu, Q. Lin, Z. Wu, Y. Li, L. Xie. 2010. from Vietnam exert suppressor activity on the RNA silencing Identification of Pns6, a putative movement protein of RRSV, pathway. Virus Genes 51(2): 267-75. doi: 10.1007/s11262- as a silencing suppressor. Virology Journal (7):335-740. 015-1229-2. Yaqoob, M., R.A. Mann, S.M. Iqbal, and M. Anwar. 2011. Reaction Nuque, F.L, Aguiero, V.M., and Ou, S.H. 1982. Inheritance of of rice genotypes to brown spot disease pathogen Cochliobolus resistance to grassy rtunt virus in rice. Plant Disease 66:63- miyabeanus under drought condition. Mycopath 9(1): 9-11. 64. Zhang, Q, 2007. Strategies for developing green super rice, PNAS 104:43, pp 16402-16409.

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