Preferred name: Xanthomonas oryzae pv. oryzicola
Authority: (Fang et al.) Swings et al.
Taxonomic position: Bacteria: Proteobacteria: Gammaproteobacteria: Lysobacterales: Lysobacteraceae
Other scientific names: Xanthomonas campestris pv. oryzicola (Fang et al.) Dye, Xanthomonas oryzicola Fang et al., Xanthomonas translucens f. sp. oryzicola (Fang et al.) Bradbury
Common names in English: BLS, bacterial leaf streak of rice, leaf streak of rice
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Notes on taxonomy and nomenclature EPPO Categorization: A1 list
EU Categorization: A1 Quarantine pest (Annex II A)
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EPPO Code: XANTTO

The principal host of X. oryzae pv. oryzicola is rice, Oryza sativa. The sticky, short-grained O. sativa subsp. Japonica (syn. O. oryza subsp. Sinica) is less susceptible to X. oryzae pv. oryzicola than the non-sticky, long-grained O. sativa subsp. Indica. In Europe, O. sativa subsp. Japonica is mainly grown (Agri-food Data Portal, 2022; Cai et al., 2013; Kraehmer et al., 2017).

Other important hosts belong to the Poaceae family, both wild and cultivated, annual and perennial species (Reddy & Nayak, 1975; Leyns et al., 1984; CABI, 2021; EFSA, 2018). 

Host list: Brachiaria lata, Digitaria horizontalis, Echinochloa colonum, Eleusine indica, Leersia hexandra, Leptochloa mucronata, Oryza barthii, Oryza glaberrima, Oryza latifolia, Oryza longistaminata, Oryza minuta, Oryza officinalis, Oryza sativa, Paspalum scrobiculatum, Paspalum vaginatum, Rottboellia cochinchinensis, Zizania aquatica, Zizania palustris, Zoysia japonica

Bacterial leaf streak was first reported in the Philippines in 1918 and is widely present in tropical and subtropical Asia, including China, Malaysia, India, Indonesia, and also in Northern Australia (under the old and incorrect name Xanthomonas translucens f.sp. oryzae, Aldrick et al., 1973) and West and East Africa, including Madagascar (CABI/EPPO, 2015). It has not been reported from temperate regions, and unlike X. oryzae pv. oryzae (see EPPO Datasheet on X. oryzae pv. oryzae) no geographically distinct groups have been determined (Ou, 1985; Mew, 1991).

Africa: Burkina Faso, Burundi, Cote d'Ivoire, Guinea, Kenya, Madagascar, Mali, Nigeria, Senegal, Uganda
Asia: Bangladesh, Cambodia, China (Anhui, Fujian, Guangdong, Guangxi, Guizhou, Hainan, Hunan, Jiangsu, Jiangxi, Sichuan, Yunnan, Zhejiang), India (Andhra Pradesh, Bihar, Haryana, Karnataka, Madhya Pradesh, Maharashtra, Uttar Pradesh, West Bengal), Indonesia (Java, Kalimantan, Sulawesi, Sumatra), Laos, Malaysia (Sabah, Sarawak, West), Myanmar, Nepal, Pakistan, Philippines, Thailand, Vietnam
Oceania: Australia (Northern Territory)

X. oryzae pv. oryzicola usually enters the host plant through stomata or leaf lesions caused by insects, heavy rain and/or wind. It multiplies in the apoplast of mesophilic parenchyma cells and spreads actively in the intercellular spaces. It causes linear water-soaked to necrotic leaf streaks, without entering the vascular tissues (Mew, 1993). X. oryzae pv. oryzicola has a strong cell-wall degrading (cellulose) activity. This differs from X. oryzae pv. oryzae which mainly infects the plant via hydathodes (water pores, connected to vascular tissue) and multiplies and spreads mainly in the vascular tissue (Tsuno & Wakimoto, 1983; Zou et al., 2012, Cao et al., 2020).

In severe infections, X. oryzae pv. oryzicola may produce typical yellow orange (amber-coloured) exudate in the form of tiny droplets from stomata on the leaf surface. The droplets dry in the form of sticky tiny beads with or without small stalks, or also in strands. These strands may be spread by dry wind (Ou, 1985; Mew, 1991). 

Both X. oryzae pv oryzicola and X. oryzae pv. oryzae can be isolated from the rice seed coat (Sakthivel et al., 2001; Niño-Liu et al., 2006), but only X. oryzae pv. oryzicola has been reported to be seed transmitted (Fang et al, 1957; Shekhawat, 1969; Mew, 1993; Xie & Mew, 1998; EFSA 2018). The bacterium can survive up to 5 months in seeds stored at 15-30 °C and seed transmission is efficient when sown under conditions of high humidity (Devadath, 1984).

The bacterium can persist from one season to the next on infected leaves and leaf debris, but was found not to survive in non-sterile soil (Devadath & Dath, 1970). The bacteria may survive on and in alternate hosts, such as Leersia hexandra and Zizania aquatica (Reddy & Nayak, 1975; Leyns et al., 1984), but this has been infrequently and/or inadequately reported (Ou, 1985; Niño-Liu et al., 2006).

Spread within a crop occurs by mechanical contact and via rain and irrigation water. Under favourable conditions (warm and wet with heavy winds) rapid and severe disease development can occur. The bacterium survives for up to 90 days in water at 15-20°C and up to 60 days at 25-45°C (Devadath, 1984). Contaminated irrigation water may spread the bacterium to adjacent fields (Devadath, 1984).

X. oryzae pv. oryzicola occurs mostly in tropical and subtropical climates and causes damage only under very wet conditions. Without continuous rain, secondary infections no longer occur (Mew, 1993; EFSA 2018).

After infection, temperature is the main determinant of disease development. Higher temperatures (26 - 32°C) favour disease development, lower temperature (below 22°C) restrain it (Devadath, 1984). Heavy nitrogenous fertilization favours disease development as is the case for X. oryzae pv. oryzae (Devadath, 1984). Insects (such as leafhoppers and grasshoppers), humans, and agricultural equipment can mechanically transmit the bacterium (Devadath, 1984). There is an apparent connection with pest damage since the bacterium readily enters insect-damaged tissue, but the exact role of these insects and that of man and machines is poorly understood. 

The pathogenicity of X. oryzae pv. oryzicola, as for X. oryzae pv. oryzae, is based on a type-3 secretion system, that injects a range of type-3 effectors into rice cells (Niño-Liu et al., 2006; Jiang et al., 2020). This includes members of the Transcription Activator-Like Effector family (TALEs), major virulence factors, activating susceptibility genes of the host (Hutin et al., 2015). Contrary to X. oryzae pv. oryzae, which has widely present gene-for-gene resistance based on an avirulence gene (bacterium) and a resistance gene (plant), so called avr-R gene interactions, this has not been identified in the X. oryzae pv. oryzicola-rice pathosystem. Resistant rice varieties therefore only show (partial), so-called quantitative resistance (Niño-Liu et al., 2006; Zhao et al., 2004; Hajri et al 2012; Cai et al, 2017). However, to date, the avrRxo1 effector gene was found to be present in all Asian X. oryzae pv. oryzicola strains, and as it is likely to be involved in fitness/pathogenicity it is therefore important for resistance breeding (Zhao et al. 2004). 

A high degree of genetic diversity was observed among Asian (Philippines) and African strains of X. oryzae pv. oryzicola. Strains from Mali were found to be closely related to those from Malaysia, implicating a possible transfer of the bacterium with planting material from Asia to Africa (Raymundo et al. 1999; Gonzalez et al., 2007; Wonni et al., 2014). In an extensive study, using 75 X. oryzae pv. oryzicola strains from South-West China and 6 differential rice varieties, Wang et al. (2010) discriminated 13 race groups, that showed some geographical differentiation. Yang et al. (2020) could discriminate 6 pathotypes of X. oryzae pv. oryzicola in Southern China, using differential varieties, and these local rice varieties showed various levels of resistance against X. oryzae pv. oryzicola.

Symptoms

Early symptoms are narrow, dark-green, water-soaked, interveinal streaks of various lengths, initially restricted to the leaf blades. The lesions enlarge, often showing a yellow halo and later turn yellowish-orange to brown (depending on the rice cultivar) and may coalesce. Bacterial ooze is often present on the streaks, visible as tiny amber-coloured drops. In advanced stages, the disease is difficult to distinguish from that caused by X. oryzae pv. oryzae but lesion margins remain linear (rather than wavy for those caused by X. oryzae pv. oryzae). It can be noted also that both X. oryzae pv. oryzicola and X. oryzae pv. oryzae may occur simultaneously in the same field, and sometimes even in the same plant (Goto, 1992; Mew, 1993). In a final stage, streaks become brown to greyish and may completely wither. Infected florets turn brown or black and the ovary and stamens die. Symptomatic infected seeds show browning of glumes and necrotic endosperm. Symptoms are often associated with those caused by larvae of lepidopterous leaf rollers/folders (e.g., Cnaphalocrocis medinalis), and of the rice hispa beetle (Discladispa armigera), because bacteria readily enter the damaged tissue resulting from these insect infestations (Ou, 1985; Niño-Liu et al., 2006; EFSA, 2018).

Morphology

X. oryzae pv. oryzicola is an aerobic, motile, Gram-negative, non-spore-forming, capsulated rod, occurring singly or in pairs, 1.0-2.5 x 0.4-0.6 µm in size, with one polar flagellum (Bradbury 1970, 1986).

Most of the procedures described for the isolation of X. oryzae pv. oryzae from rice plants, can also be applied for the isolation of X. oryzae pv. oryzicola. (EPPO, 2007). Faster growing contaminants often occurring on and in diseased tissues, such as species of Pantoea or xanthomonad-like saprophytes may overgrow the slow growing X. oryzae pv. oryzicola colonies and hinder its isolation from diseased material. 

Isolation of X. oryzae pv. oryzicola from symptomatic material is possible on Peptone Sucrose Agar (PSA), Nutrient Broth Yeast Extract agar medium (NBY), Growth Factor (GF) agar or otherwise using semi- selective media (Agarwal et al., 1989; Sakthivel et al.,2001; EPPO, 2007). A semi-selective medium, called XOS, is available for detection of X. oryzae pv. oryzicola from rice seed (Di et al., 1991; EPPO, 2007). On nutrient agar (NA), after 3 days of growth, colonies of X. oryzae pv. oryzicola are circular, entire, smooth, convex, opaque, and pale to straw yellow, 1-2 mm in size. Optimum growth temperature is between 25 and 30°C. For growth on other media, see EPPO, 2007. 

Detection and identification methods

Detection of X. oryzae pv. oryzicola in seed, using a detached leaf inoculation method was described by Xie & Mew (1998). The method is based on inoculating leaf segments on agar with seed washings in a moist chamber. For selective recovery from seed, this method and the XOS semi-selective medium of Di et al. (1991) can be used. 

Furthermore poly- and monoclonal antibodies (genus and pathovar specific) can be used in Immuno-fluorescence and ELISA tests on seed extracts and/or colonies isolated from seeds or leaf/stem material and isolated bacterial cells (Benedict et al., 1989). An ELISA kit is commercially available for the detection of X. oryzae pv. oryzicola (EPPO, 2007). A padlock probe (PLP)-based PCR with dot blot hybridisation was developed for simultaneous detection of X. oryzae pv. oryzicola and X. oryzae pv. oryzae by Tian et al., 2014. A specific TaqMan probe for its detection in seed was developed by Zhao et al. (2007).

Leach et al. (1990) used a repetitive DNA sequence (pJEL 101) to distinguish X. oryzae pv. oryzae from other pathovars and species of Xanthomonas. Kang et al. (2008) developed a specific PCR detection system (targeting a membrane fusion protein gene) for X. oryzae pv. oryzicola. Other specific TaqMan-based multiplex PCRs for detection and discrimination of X. oryzae pv. oryzicola and X. oryzae pv. oryzae were developed and validated by Lang et al. (2010), Noh et al. (2012), Kang et al (2012) and Lee & Vera Cruz (2014). 

Lang et al. (2014) developed a sensitive and rapid loop-mediated isothermal amplification (LAMP) test, using primer sets to distinguish not only X. oryzae pv. oryzicola and X. oryzae pv. oryzae, but also the Asian and African lines of X. oryzae pv. oryzae.

A SYBR green-based multiplex PCR for the detection and identification of X. oryzae pv. oryzicola, X. oryzae pv. oryzae and Burkholderia glumae (causing bacterial grain rot of rice) was developed by Lu et al. (2014). Kang et al. (2016) also developed a multiplex PCR for the detection of the same three bacteria. Cui et al. (2016) developed a multiplex conventional and real-time PCR for the simultaneous detection of six bacterial pathogens of rice, including X. oryzae pv. oryzicola, X. oryzae pv. oryzae, Pseudomonas fuscovaginae (rice sheath brown rot), Burkholderia glumae, B. gladioli (bacterial panicle blight of rice) and Acidovorax avenae subsp. avenae (bacterial brown stripe of rice). A validated multiplex PCR to detect P. fuscovaginae, X. oryzae pv. oryzicola and X. oryzae pv. oryzae, Burkholderia (both B. glumae and B. gladioli) as well as Sphingomonas and Pantoea spp. was published by Bangratz et al. (2020).

The two pathovars of X. oryzae differ in the symptoms induced (Ou, 1985), phenotypic characters (Reddy & Ou, 1974; Vera Cruz et al., 1984; Vauterin et al., 1995), polyacrylamide gel electrophoresis protein fingerprints (Mew & Vera Cruz, 1979; Kersters et al., 1989), serological behavior (Benedict et al., 1989) and phage typing (EPPO 2007). Also, on the basis of rep-PCR using BOX-primers discrimination of X. oryzae pv. oryzicola and X. oryzae pv. oryzae is possible (Raymundo et al., 2008).

As for the whole genus Xanthomonas, X. oryzae is catalase-positive, unable to reduce nitrate and a weak producer of acids from carbohydrates. Pathovars oryzicola and oryzae can be differentiated by (a) acetoin production (X. oryzae pv. oryzicola+, X. oryzae pv. oryzae–), (b) growth on l-alanine as sole carbon source (X. oryzae pv. oryzicola+, X. oryzae pv. oryzae–), (c) growth on 0.2% vitamin-free casamino acids (X. oryzae pv. oryzicola+, X. oryzae pv. oryzae–) and (d) resistance to 0.001% Cu (NO₃)₂ (X. oryzae pv. oryzicola–, X. oryzae pv. oryzae+) (Dye & Lelliott, 1974; Reddy & Ou, 1974; Gossele et al., 1985; Niño-Liu et al., 2006; EPPO 2007). Extensive characterization of X. oryzae pv. oryzicola, using biochemical, physiological tests and PAGE was performed by Vera Cruz et al. (1984). Wonni et al. (2014) determined extensive variability between African strains of X. oryzae pv. oryzicola. Restriction fragment length polymorphism (RFLP) analysis using the effector avrXa7 as probe resulted in the identification of 18 haplotypes. PCR using two conserved type III effector (T3E) genes (xopAJ and xopW) differentiated the strains into an African group where the xopAJ was generally not detected, and a group of possible Asian origin. 

Six housekeeping genes— atpD (ATP synthase β chain), dnaK (chaperone protein), efP (elongation factor P), gyrB (DNA gyrase subunit B), lepA (GTP binding protein), and especially rpoD (RNA polymerase σ-70 factor) are useful for identification and phylogenetic studies of X. oryzae pv. oryzicola strains (Afolabi et al., 2014; Wonni et al., 2014).

Isolates can be tested for pathogenicity on susceptible rice cultivars. For X. oryzae pv. oryzicola 30–45-day old plants of cultivars IR24 or IR50 (International Rice Institute) or local, susceptible varieties can be used. Leaf clipping and spray inoculation methods are available for inoculations (Kauffman et al., 1973; Cottyn et al., 1994; EPPO, 2007; Afolabi et al., 2014). Niño-Liu et al. (2005) inoculated plants by dipping them in bacterial mixture and incubating in a growth chamber. Symptoms developed within 6 days.

X. oryzae pv. oryzicola can only move short distances within infected crops. The bacterium is found in association with weeds, even if their role in the disease cycle is less clear than for X. oryzae pv. oryzae (Leyns et al., 1984; Reddy & Nayak, 1975).

There are little substantiated data on spread or transmission in the field by animals other than insects (Ou, 1985; Niño-Liu et al., 2006; EFSA, 2018).

Long distance spread can take place via infected rice seeds, and seed transmission is regarded as the main means of dispersal. The planting of disease-free seed is considered of utmost importance in control (Rao, 1987; Xie et al., 1990, 1991; Ming et al., 1991; Mew, 1993; Xie & Mew, 1998).

Economic impact

Bacterial leaf streak is only of importance in some areas during very wet seasons and where high levels of nitrogen fertilization are used. It does not usually reduce yields if low levels of nitrogen fertilization are applied. In general, bacterial leaf streak is a much less important disease than bacterial leaf blight. In Central India, losses ranged from 5 to 30% depending upon environmental factors and cultivars (Naik et al., 1973). In Northern India, disease intensity affecting 80% of leaf area resulted in 61 percent yield loss (Singh et al., 1980). In the Philippines, no significant losses were reported in either the wet or dry seasons (Opina & Exconde, 1971).

In West Africa outbreaks of X. oryzae pv. oryzicola usually showed lower incidence and severity than those of X. oryzae pv. oryzae, as determined in a 10-year survey (Awoderv et al., 1991).

In China, however, X. oryzae pv. oryzicola has sometimes been more damaging than X. oryzae pv. oryzae. In Southern China, epidemics of X. oryzae pv. oryzicola have repeatedly been reported, reducing yield by 10-20% and in some cases reaching up to 40% losses (Xie & Mew 1998; Niño-Liu et al., 2006; Cai et al., 2017). In Uganda, under favourable conditions (wet/windy/warm temperatures/susceptible varieties) bacterial leaf streak has caused major crop losses (up to 60%) (Andaku et al., 2016; EFSA, 2018).

Control

The bacterial leaf streak pathogen hardly requires any particular control measures except the use of healthy seed and prevention measures (see below). Neither treatments nor resistance are mentioned to any significant extent in the literature.

Chemical control

Chemical seed treatment and field sprays have been reported from India (Shekhawat & Srivastava 1971), using a combination of antibiotics (streptomycin sulphate and tetracycline) and copper-oxychloride. It was also reported that when yield is affected, a copper-based fungicide applied at heading stage can be effective in controlling the disease (ICAR/TNAU, 2022; CABI Plantwise, 2022). The use of antibiotics against plant pathogens is not permitted in many EPPO countries, although in Asia their use is still ongoing and resistance against streptomycin has been reported in China (Xu et al., 2010). Recently Chen et al. (2019) reported a strong bactericidal effect (in vitro and in vivo) of the bactericide melatonin (N-acetyl-5-methoxytryptamine) on X. oryzae pv. oryzicola and a reduction of disease incidence by 17%.

Heat treatment

Hot water treatment of rice seeds at 52-54°C for 30 min, preceded by 8-10 hour of presoaking at room temperature in water, has been advised and used to cure seeds of X. oryzae pv. oryzae (Jain, 1970; Reddy, 1983) and is also expected to be effective for X. oryzae pv. oryzicola. 

Biological control

Hata et al. (2015) found an antagonistic effect on X. oryzae pv. oryzicola of Streptomyces spp. in vitro. In a follow-up (greenhouse) study two strains showed a suppressive effect on bacterial leaf streak due to induction of systemic resistance and growth promoting activity (Hata et al., 2021)

Zhang et al. (2012) reported promising biocontrol effect of strain Lx-11 of Bacillus amyloliquefaciens. This strain appears also to trigger a systemic immunization activity and significantly reduced disease incidence in field experiments (from 60% to 71%) which was better than the effect of a chemical spray with thiadiazole-copper (a bactericide often used in China).

Plant resistance

In contrast to bacterial leaf blight, native major resistance genes controlling resistance to bacterial leaf streak have not yet been identified in rice. There are, however loci determining quantitative resistance, such as qBLSR-11-1 (Chen et al., 2006) and qBlsr5a, which had a relatively large impact in breeding lines, where the broadly effective rice recessive gene xa5 is involved (Xie et al., 2020). In a genome-wide resistance-gene analysis in rice Sattayachiti et al. (2020) and Thianthavon et al. (2021) stated that this recessive xa5 gene is a very promising candidate to be used in breeding for broad-spectrum resistance. A non-host resistance gene, Rxo1, isolated from maize, and present in transgenic rice was shown to confer high level resistance to bacterial leaf streak (Zhao et al., 2005; Jiang et al., 2020). Using CRISPR/Cas9 gene editing of two rice varieties Ni et al. (2021) obtained rice lines that proved to be resistant to X. oryzae pv. oryzicola and X. oryzae pv. oryzae. The original agronomic traits of these lines were not diminished. The dominant locus Xo1 apparently confers complete resistance to African strains of X. oryzae pv. oryzicola (Triplett et al., 2016, Cai et al., 2017).

Prevention and cultural control

Prophylactic measures (such as use of healthy seeds, adequate fertilization and irrigation, destruction or ploughing under of crop residues, disinfection of machinery and equipment, production of seedlings in boxes and removal of diseased plants and weed hosts from fields and along irrigation canals) have all been found useful in the control of bacterial leaf streak (Devadath, 1984; Goto, 1992; Ou 1985; Shekhawat et al., 1972).

Phytosanitary risk

Rice cultivation in Europe occurs in Bulgaria, France, Greece, Hungary, Italy Portugal, Romania, the Russian Federation, Spain, Turkey and Ukraine. About 80% of the European Union rice production takes place in Italy (>220 000 ha and Spain (>115 000 ha), another 12% in Greece and Portugal (some 20-25 000 ha each). The remainder is cultivated in Bulgaria, France, Hungary and Romania, (10-20 000 ha each). In non-EU European countries, rice is grown in the Russian Federation (120 000 ha in the Krasnodar region) as well as in Ukraine (25 000 ha). In those countries all rice fields are under irrigation, planted in spring and harvested in autumn. (Agri-food Data Portal, 2022; Ferrero & Nguyen, 2004; Kraehmer et al., 2017).

Resistance of European varieties against X. oryzae pv. oryzicola is unknown. Non-European varieties are only introduced, in small quantities, for breeding (Cai et al., 2013; Kraehmer et al., 2017). No interceptions of X. oryzae pv oryzicola have been reported in the EU from 1995 to April 2022 (European Commission, 2022). However, no systematic surveying and monitoring for X. oryzae pv. oryzicola takes place in Europe

The main risk of introduction is via imported rice seed (germplasm) used for breeding purposes and therefore direct sowing. Milled rice poses a negligible risk, because hulls are removed, and endosperm infection is very rare. Moreover, milled rice has its main destination outside growing areas. 

Once introduced by infected seed, further spread could take place via newly infected seed and contaminated water and the bacterium could survive in stubble, straw, weed hosts and volunteer plants.

Phytosanitary (quarantine) measures can be implemented to reduce the risk of long-distance dissemination of the pathogen. It can be recommended that consignments of rice seeds should have been produced from pest-free areas, or from pest-free places of production.

General inspection and sampling procedures for imported rice, which include X. oryzae pv. oryzicola are described in EPPO Standard PM 3/78(2) Consignment inspection of seed and grain of cereals. Seed inspections of rice intended for breeding purposes in international trade may assist in preventing spread of the pathogen to areas with no history of the disease. However, visual inspection of imported seeds is not very reliable due to the occurrence of latent infections and therefore, when material is imported from areas where the disease is known to occur, certification for disease freedom via field inspections and laboratory testing are necessary. 

A contingency plan to prepare for possible introductions of X. oryzae pv. oryzicola in the USA, was developed by the USDA (USDA, 2013).

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CABI and EFSA resources used when preparing this datasheet

CABI (2022) Xanthomonas oryzae pv. oryzae (bacterial leaf blight of rice). https://www.cabi.org/isc/datasheet/56956 [Accessed: April 2022].

CABI (2022) Xanthomonas oryzae pv. oryzicola (bacterial leaf streak of rice). https://www.cabi.org/isc/datasheet/56977 [Accessed: April 2022].

EFSA (2018) EFSA Panel on Plant Health: Jeger M, Candresse T, Chatzivassiliou E, Dehnen-Schmutz K, Gilioli G, Gregoire J-C, Jaques Miret JA, MacLeod A, Navajas Navarro M, Niere B, Parnell S, Potting R, Rafoss T, Rossi V, Urek G, Van Bruggen A, Van der Werf W, West J, Winter S, Bragard C, Szurek B, Hollo G and Caffier D. Scientific Opinion on the pest categorisation of Xanthomonas oryzae pathovars oryzae and oryzicola. EFSA Journal 16, 5109, 25 pp. https://doi.org/10.2903/j.efsa.2018.5109

This datasheet was extensively revised in 2022 by Dr Jaap D. Janse, independent consultant, bacteriologist. His valuable contribution is gratefully acknowledged.

EPPO (2024) Xanthomonas oryzae pv. oryzicola. EPPO datasheets on pests recommended for regulation. https://gd.eppo.int (accessed 2024-12-25)

This datasheet was first published in the EPPO Bulletin in 1980 (as Xanthomonas oryzae) and revised in the two editions of 'Quarantine Pests for Europe' in 1992 and 1997, as well as in 2022. It is now maintained in an electronic format in the EPPO Global Database. The sections on 'Identity', ‘Hosts’, and 'Geographical distribution' are automatically updated from the database. For other sections, the date of last revision is indicated on the right.

CABI/EPPO (1992/1997) Quarantine Pests for Europe (1st and 2nd edition). CABI, Wallingford (GB).

EPPO (1980) Data sheets on quarantine organisms No. 2, Xanthomonas oryzae. EPPO Bulletin 10(1), 4 pp. https://doi.org/10.1111/j.1365-2338.1980.tb02685.x