Aphelenchoides besseyi (rice leaf nematode)

Pictures

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Picture	Title	Caption	Copyright
Title Symptoms on leaves Caption Left: Characteristic 'white tip' symptom on rice leaf. Right: Necrotic patches and crinkled rice leaves. Copyright John Bridge/CABI BioScience	Symptoms on leaves	Left: Characteristic 'white tip' symptom on rice leaf. Right: Necrotic patches and crinkled rice leaves.	John Bridge/CABI BioScience

Identity

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Preferred Scientific Name

• Aphelenchoides besseyi Christie 1942

Preferred Common Name

• rice leaf nematode

Other Scientific Names

• Aphelenchoides oryzae Yokoo 1948
• Asteroaphelenchoides besseyi (Christie 1942) Drozdovski 1967

International Common Names

• English: summer crimp nematode; white tip; white tip nematode of rice
• Spanish: nematodo de la punta blanca del arroz
• French: nématode du bout blanc du riz; nématode foliaire du riz

EPPO code

• APLOBE (Aphelenchoides besseyi)

Taxonomic Tree

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• Domain: Eukaryota
•     Kingdom: Metazoa
•         Phylum: Nematoda
•             Order: Aphelenchida
•                 Family: Aphelenchoididae
•                     Genus: Aphelenchoides
•                         Species: Aphelenchoides besseyi

Description

Top of page Measurements

(from type host and locality after Christie, 1942):

* 10 females: L = 0.66-0.75 mm; a = 32-42 (width = 17-22 µm): b = 10.2-11.4 (oesophagus = 64-68 µm): c = 17-21 (tail = 36-42 µm ); V = 68-70.

* 10 males: L = 0.54-0.62 mm; a = 36-39 (width = 14-17 µm); b = 8.6-8.8 (oesophagus = 63-66 µm); c = 15-17 (tail = 34-37 µm).


(After Allen, 1952):

* females: L = 0.62-0.88 mm; a = 38-58; b = 9-12; c = 15-20; V = 43-3366-724-8.

* males: L = 0.44-0.72 mm; a = 36-47; b = 9-11; c = 14-19; T = 50-65.


(From rice seeds from Séfa, Senegal; after Fortuner, 1970):

* 20 females: L = 0.57-0.84 (0.68) mm; a = 39-53 (47.7); b = 9.2-13.1 (11.46); b' = 4.06-5.77 (4.85); c = 13.8-20.4 (17.7); V = 39..1-19..968.7-73.6 (71.2)4.1-6.2; spear = 10.0-12.5 (11.9) µm.

* 9 males: L = 0.53-0.61 (0.57) mm; a = 40.7-46.9 (44.4); b = 8.87-10.70 (9.52); b' = 3.57-4.91 (4.09); c = 16-20 (17.97); T = 28-52 (40.59); spear = 10.0-12.5 (11.4) µm; spicules (dorsal limb) = 18-21 (19.2) µm.


Description (after Franklin & Siddiqi, 1972)

Female
The body of female A. besseyi is slender, straight to slightly arcuate ventrally when relaxed; annules fine, indistinct, about 0.9 µm wide near mid-body. Lip region rounded, unstriated, slightly offset and wider than body at lip base, about half as wide as mid-body; labial framework hexaradiate, lightly sclerotized. Lateral fields about one-fourth as wide as body, with 4 incisures. Anterior part of spear sharply pointed, about 45% of total spear length, posterior part with slight basal swellings which are 1.75 µm across. Median oesophageal bulb oval, with a distinct valvular apparatus slightly behind its centre. Oesophageal glands extending dorsally and subdorsally for 4 to 8 body-widths over intestine. Nerve ring about one body-width behind median oeosphageal bulb.

Excretory pore usually near anterior edge of nerve ring. Hemizonid 11-15 µm behind excretory pore. Vulva transverse, with slightly raised lips. Spermatheca elongate oval (up to 8 times as long as wide when fully distended), usually packed with sperm. Ovary relatively short and not extending to oesophageal glands, with oocytes in 2-4 rows. Post-vulva uterine sac narrow, inconspicuous, not containing sperm, 2.5-3.5 times anal body width long but less than one-third distance from vulva to anus. Tail conoid, 3.5-5 anal body widths long; terminus bearing a mucro of diverse shape with 3-4 pointed processes.

Male
Male A. besseyi are about as numerous as females. The posterior end of body is curved to about 180 degrees in relaxed specimens. Lip region, spear and oesophagus as described for female; tail conoid, with terminal mucro with 2-4 pointed processes. First pair of ventrosubmedian papillae adanal, second slightly behind middle of tail and third subterminal. Spicules typical of the genus except that the proximal end lacks a dorsal process (apex) and has only a moderately developed ventral one (rostrum). Testis single, oustretched.

Distribution

Top of page A. besseyi is very widely distributed and now occurs in most rice growing areas (Ou, 1985). Its wide distribution has resulted from dissemination in seed.


See also CABI/EPPO (1998, No. 157).

Distribution Table

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The distribution in this summary table is based on all the information available. When several references are cited, they may give conflicting information on the status. Further details may be available for individual references in the Distribution Table Details section which can be selected by going to Generate Report.

Continent/Country/Region	Distribution	Last Reported	Origin	First Reported	Invasive	Reference	Notes
Asia
Afghanistan	Present					CABI/EPPO, 2000; EPPO, 2014
Azerbaijan	Present					CABI/EPPO, 2000; EPPO, 2014
Bangladesh	Widespread					Timm, 1955; CABI/EPPO, 2000; EPPO, 2014
Cambodia	Present					CABI/EPPO, 2000; EPPO, 2014
China	Present					Qiu et al., 1991; CABI/EPPO, 2000; EPPO, 2014
-Anhui	Present					CABI/EPPO, 2000; EPPO, 2014
-Fujian	Present					CABI/EPPO, 2000; EPPO, 2014
-Guangdong	Present					CABI/EPPO, 2000; EPPO, 2014
-Guangxi	Present					CABI/EPPO, 2000; EPPO, 2014
-Guizhou	Present					CABI/EPPO, 2000; EPPO, 2014
-Hebei	Present					CABI/EPPO, 2000; EPPO, 2014
-Henan	Present					CABI/EPPO, 2000; EPPO, 2014
-Hubei	Present					CABI/EPPO, 2000; EPPO, 2014
-Hunan	Present					CABI/EPPO, 2000; EPPO, 2014
-Jiangsu	Present					CABI/EPPO, 2000; EPPO, 2014
-Jiangxi	Present					CABI/EPPO, 2000; EPPO, 2014
-Jilin	Present					Ou et al., 2014
-Liaoning	Present					CABI/EPPO, 2000; EPPO, 2014
-Shaanxi	Present					CABI/EPPO, 2000; EPPO, 2014
-Shandong	Present					CABI/EPPO, 2000; EPPO, 2014
-Shanxi	Present					CABI/EPPO, 2000; EPPO, 2014
-Sichuan	Present					CABI/EPPO, 2000; EPPO, 2014
-Xinjiang	Present					CABI/EPPO, 2000; EPPO, 2014
-Zhejiang	Present					CABI/EPPO, 2000; EPPO, 2014
Georgia (Republic of)	Widespread					CABI/EPPO, 2000; EPPO, 2014
India	Widespread					Prasad et al., 1987; CABI/EPPO, 2000; EPPO, 2014
-Andaman and Nicobar Islands	Present					Nandini et al., 2001
-Andhra Pradesh	Present					Prasad et al., 1987; CABI/EPPO, 2000; EPPO, 2014
-Assam	Widespread					Prasad et al., 1987; CABI/EPPO, 2000; EPPO, 2014
-Bihar	Widespread					Prasad et al., 1987; CABI/EPPO, 2000; EPPO, 2014
-Delhi	Widespread					Prasad et al., 1987; CABI/EPPO, 2000; EPPO, 2014
-Gujarat	Present					Prasad et al., 1987; CABI/EPPO, 2000; EPPO, 2014
-Haryana	Widespread					Prasad et al., 1987; CABI/EPPO, 2000; EPPO, 2014
-Indian Punjab	Present					CABI/EPPO, 2000; EPPO, 2014
-Jammu and Kashmir	Present					Munshi et al., 2001
-Karnataka	Present					Ramakrishnan and Rajendran, 2003
-Kerala	Present					Prasad et al., 1987; CABI/EPPO, 2000; EPPO, 2014
-Madhya Pradesh	Widespread					Prasad et al., 1987; CABI/EPPO, 2000; EPPO, 2014
-Maharashtra	Widespread					Prasad et al., 1987; CABI/EPPO, 2000; EPPO, 2014
-Meghalaya	Present					CABI/EPPO, 2000; EPPO, 2014
-Odisha	Present					Prasad et al., 1987; CABI/EPPO, 2000; EPPO, 2014
-Tamil Nadu	Present					Prasad et al., 1987; CABI/EPPO, 2000; EPPO, 2014
-Tripura	Widespread					Prasad et al., 1987; CABI/EPPO, 2000; EPPO, 2014
-Uttar Pradesh	Widespread					Prasad et al., 1987; CABI/EPPO, 2000; EPPO, 2014
-West Bengal	Present					Prasad et al., 1987; CABI/EPPO, 2000; EPPO, 2014
Indonesia	Present					Anon., 1981; CABI/EPPO, 2000; EPPO, 2014
Iran	Present					CABI/EPPO, 2000; EPPO, 2014
Israel	Absent, formerly present					CABI/EPPO, 2000; EPPO, 2014
Japan	Present					Kobayashi, 1976; CABI/EPPO, 2000; EPPO, 2014
-Honshu	Present					CABI/EPPO, 2000; EPPO, 2014
-Kyushu	Present					CABI/EPPO, 2000; EPPO, 2014
-Shikoku	Present					CABI/EPPO, 2000; EPPO, 2014
Korea, DPR	Present					Choo and Choi, 1979
Korea, Republic of	Present					Kim and Han, 1989; CABI/EPPO, 2000; EPPO, 2014
Kyrgyzstan	Present					CABI/EPPO, 2000; EPPO, 2014
Laos	Present					CABI/EPPO, 2000; EPPO, 2014
Malaysia	Present					Muhammed, 1992; CABI/EPPO, 2000; EPPO, 2014
-Peninsular Malaysia	Present					Rahim, 1988; CABI/EPPO, 2000; EPPO, 2014
Myanmar	Present					CABI/EPPO, 2000; EPPO, 2014
Nepal	Restricted distribution					CABI/EPPO, 2000; EPPO, 2014
Pakistan	Present					CABI/EPPO, 2000; EPPO, 2014
Philippines	Present					Timm, 1965; CABI/EPPO, 2000; EPPO, 2014
Singapore	Present, few occurrences					AVA, 2001
Sri Lanka	Present					CABI/EPPO, 2000; EPPO, 2014
Taiwan	Widespread					Wang et al., 1993; CABI/EPPO, 2000; EPPO, 2014
Tajikistan	Restricted distribution					CABI/EPPO, 2000; EPPO, 2014
Thailand	Present					Timm, 1965; CABI/EPPO, 2000; EPPO, 2014
Turkey	Restricted distribution					CABI/EPPO, 2000; EPPO, 2014
Uzbekistan	Present					CABI/EPPO, 2000; EPPO, 2014
Vietnam	Present					Sudakova et al., 1964; CABI/EPPO, 2000; EPPO, 2014
Africa
Benin	Present					Barat et al., 1969; CABI/EPPO, 2000; EPPO, 2014
Burkina Faso	Present					Fortuner, 1970; CABI/EPPO, 2000; EPPO, 2014
Burundi	Present					CABI/EPPO, 2000; EPPO, 2014
Cameroon	Present					Barat et al., 1969; CABI/EPPO, 2000; EPPO, 2014
Central African Republic	Present					Barat et al., 1969; CABI/EPPO, 2000; EPPO, 2014
Chad	Present					Barat et al., 1969; CABI/EPPO, 2000; EPPO, 2014
Comoros	Present					CABI/EPPO, 2000; EPPO, 2014
Congo Democratic Republic	Present					Barat et al., 1969; CABI/EPPO, 2000; EPPO, 2014
Côte d'Ivoire	Present					Merny, 1970; CABI/EPPO, 2000; EPPO, 2014
Egypt	Present					Barat et al., 1966; CABI/EPPO, 2000; EPPO, 2014
Gabon	Present					Barat et al., 1969; CABI/EPPO, 2000; EPPO, 2014
Gambia	Present					CABI/EPPO, 2000; EPPO, 2014
Ghana	Present					Peachey et al., 1966; CABI/EPPO, 2000; EPPO, 2014
Guinea	Present					CABI/EPPO, 2000; EPPO, 2014
Kenya	Widespread					Taylor, 1972; CABI/EPPO, 2000; EPPO, 2014
Madagascar	Present					Barat et al., 1969; CABI/EPPO, 2000; EPPO, 2014
Malawi	Present					Siddiqi, 1970; CABI/EPPO, 2000; EPPO, 2014
Mali	Present					Fortuner, 1970; CABI/EPPO, 2000; EPPO, 2014
Nigeria	Present					Peachey et al., 1966; CABI/EPPO, 2000; EPPO, 2014
Senegal	Present					Hooper and Merny, 1966; CABI/EPPO, 2000; EPPO, 2014
Sierra Leone	Present					Hooper and Merny, 1966; CABI/EPPO, 2000; EPPO, 2014
South Africa	Restricted distribution					CABI/EPPO, 2000; EPPO, 2014
Tanzania	Widespread					Taylor et al., 1972; CABI/EPPO, 2000; EPPO, 2014
Togo	Present					Barat et al., 1969; CABI/EPPO, 2000; EPPO, 2014
Uganda	Present					Taylor, 1972; CABI/EPPO, 2000; EPPO, 2014
Zambia	Present					CABI/EPPO, 2000; EPPO, 2014
Zimbabwe	Present					Anon., 1972; CABI/EPPO, 2000; EPPO, 2014
North America
Mexico	Present					Preciado, 1953; CABI/EPPO, 2000; EPPO, 2014
USA	Restricted distribution					Atkins and Todd, 1959; CABI/EPPO, 2000; EPPO, 2014
-Arkansas	Present					Atkins and Todd, 1959; CABI/EPPO, 2000; EPPO, 2014
-California	Present					CABI/EPPO, 2000; EPPO, 2014
-Florida	Present					CABI/EPPO, 2000; EPPO, 2014
-Hawaii	Present					Allen, 1952; CABI/EPPO, 2000; EPPO, 2014
-Louisiana	Present					Atkins and Todd, 1959; CABI/EPPO, 2000; EPPO, 2014
-Texas	Present					Atkins and Todd, 1959; CABI/EPPO, 2000; EPPO, 2014
Central America and Caribbean
Cuba	Present					Fernandez, 1967; CABI/EPPO, 2000; EPPO, 2014
Dominica	Present					CABI/EPPO, 2000; EPPO, 2014
Dominican Republic	Present					CABI/EPPO, 2000; EPPO, 2014
El Salvador	Present					Ancalmo and Davis, 1962; CABI/EPPO, 2000; EPPO, 2014
Guadeloupe	Present					Kermarrec and Anais, 1973; CABI/EPPO, 2000; EPPO, 2014
Panama	Present					CABI/EPPO, 2000; EPPO, 2014
Trinidad and Tobago	Absent, intercepted only					EPPO, 2014
South America
Argentina	Absent, invalid record					CABI/EPPO, 2000; EPPO, 2014
Brazil	Present					Lordello, 1969; CABI/EPPO, 2000; EPPO, 2014
-Bahia	Present					CABI/EPPO, 2000; EPPO, 2014
-Minas Gerais	Present					CABI/EPPO, 2000; EPPO, 2014
-Parana	Restricted distribution					CABI/EPPO, 2000; EPPO, 2014
-Rio Grande do Norte	Present					CABI/EPPO, 2000; EPPO, 2014
-Rio Grande do Sul	Present					CABI/EPPO, 2000; EPPO, 2014
-Santa Catarina	Present					CABI/EPPO, 2000; EPPO, 2014
-Sao Paulo	Present					CABI/EPPO, 2000; EPPO, 2014
Ecuador	Present, few occurrences					CABI/EPPO, 2000; EPPO, 2014
Europe
Belgium	Absent, no pest record					EPPO, 2014
Bulgaria	Restricted distribution			197*		Stoyanov, 1979; CABI/EPPO, 2000; EPPO, 2014
France	Absent, formerly present					Anon., 1981; CABI/EPPO, 2000; EPPO, 2014
Hungary	Restricted distribution			1970		Javor, 1970; CABI/EPPO, 2000; EPPO, 2014
Italy	Restricted distribution			1973		Orsenigo, 1954; CABI/EPPO, 2000; EPPO, 2014
Netherlands	Transient: actionable, under surveillance					CABI/EPPO, 2000; EPPO, 2014	Oryza: absent, no pest records; Fragaria: absent, confirmed by survey; few ornamental greenhouse crops (not related to Fragaria and Oryza): transient, actionable, under surveillance. Based on long-term annual surveys, 10 survey observations in 2012.
Portugal	Absent, confirmed by survey					EPPO, 2014
Russian Federation	Restricted distribution					CABI/EPPO, 2000; EPPO, 2014
-Southern Russia	Present					CABI/EPPO, 2000; EPPO, 2014
Slovakia	Absent, invalid record					CABI/EPPO, 2000; EPPO, 2014
UK	Absent, intercepted only					CABI/EPPO, 2000; EPPO, 2014
Ukraine	Restricted distribution					CABI/EPPO, 2000; EPPO, 2014
Oceania
Australia	Restricted distribution					CABI/EPPO, 2000; EPPO, 2014
-Australian Northern Territory	Present					CABI/EPPO, 2000; EPPO, 2014
-Queensland	Present					CABI/EPPO, 2000; EPPO, 2014
Cook Islands	Present					CABI/EPPO, 2000; EPPO, 2014
Fiji	Present					Kirby et al., 1977; CABI/EPPO, 2000; EPPO, 2014
Papua New Guinea	Present					CABI/EPPO, 2000; EPPO, 2014

Risk of Introduction

Top of page RISK CRITERIA CATEGORY

ECONOMIC IMPORTANCE High
DISTRIBUTION Worldwide
SEEDBORNE INCIDENCE High
SEED TRANSMITTED Yes
SEED TREATMENT Yes

OVERALL RISK High


Notes on phytosanitary risk

A. besseyi already has a world wide distribution but should remain a quarantine pest in rice growing countries because of the potential danger of the emergence of more virulent pathotypes. Seeds for export should be treated with hot water, unless nematode-free seed can be ensured.

Habitat

Top of page A. besseyi is able to infect rice in all rice production environments. Infection and damage are generally greater in lowland and deep water systems than in upland environments.

Host Plants and Other Plants Affected

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Plant name	Family	Context
Allium cepa (onion)	Liliaceae	Other
Begonia	Begoniaceae	Other
Chrysanthemum morifolium (chrysanthemum (florists'))	Asteraceae	Other
Colocasia esculenta (taro)	Araceae	Other
Cyperus iria (rice flatsedge)	Cyperaceae	Other
Digitaria sanguinalis (large crabgrass)	Poaceae	Other
Dioscorea (yam)	Dioscoreaceae	Other
Dioscorea trifida (cushcush yam)	Dioscoreaceae	Other
Fragaria ananassa (strawberry)	Rosaceae	Main
Glycine max (soyabean)	Fabaceae	Other
Hibiscus (rosemallows)	Malvaceae	Other
Ipomoea batatas (sweet potato)	Convolvulaceae	Other
Oryza (rice (generic level))	Poaceae	Main
Oryza breviligulata	Poaceae	Wild host
Oryza glaberrima (African rice)	Poaceae	Main
Oryza sativa (rice)	Poaceae	Main
Polianthes tuberosa (tuberose)	Agavaceae	Other
Setaria viridis (green foxtail)	Poaceae	Wild host
Zea mays (maize)	Poaceae	Other

Growth Stages

Top of page Seedling stage, Vegetative growing stage

Symptoms

Top of page Rice

Plants susceptible to A. besseyi can be symptomless but yield loss only occurs in plants showing some symptoms. During early growth, the most conspicuous symptom is the emergence of the chlorotic tips of new leaves from the leaf sheath. These tips later dry and curl, whilst the rest of the leaf may appear normal. The young leaves of infected tillers can be speckled with a white splash pattern or have distinct chlorotic areas. Leaf margins may be distorted and wrinkled but leaf sheaths are symptomless.

The viability of A. besseyi infected seed is lowered and germination is delayed (Tamura and Kegasawa, 1959b) and diseased plants have reduced vigour and height (Todd and Atkins, 1958). Infected panicles are shorter, with fewer spikelets and a smaller proportion of filled grain (Dastur, 1936; Yoshii, 1951; Todd and Atkins, 1958).

In severe infections, the shortened flagleaf is twisted and can prevent the complete extrusion of the panicle from the boot (Yoshii and Yamamoto, 1950a; Todd and Atkins, 1958). The grain is small and distorted (Todd and Atkins, 1958) and the kernel may be discoloured and cracked (Uebayashi et al., 1976). Infected plants mature late and have sterile panicles borne on tillers produced from high nodes.

Strawberry

A. besseyi is a foliar pest of strawberry and may be found between leaves and buds. Aphelenchoides spp. cause distortion of the leaves which is more noticeable on newly formed leaves after growth resumes in spring in areas of the USA, south of Virginia and Arkansas, and also in Australia (Brown et al., 1993).

List of Symptoms/Signs

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Sign	Life Stages	Type
Inflorescence / discoloration panicle
Inflorescence / twisting and distortion
Leaves / abnormal colours
Leaves / abnormal forms
Leaves / abnormal patterns
Seeds / discolorations
Seeds / empty grains
Seeds / lesions on seeds
Stems / stunting or rosetting
Whole plant / dwarfing

Biology and Ecology

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Rice

After sowing, anabiotic A. besseyi rapidly become active and are attracted to meristematic areas. During early growth, it is found in low numbers within the innermost leaf sheath, feeding ectoparasitically around the apical meristem (Yoshii and Yamamoto, 1950b; Goto and Fukatsu, 1952; Todd and Atkins, 1958). The main stem is frequently more infected than subsequent tillers (Goto and Fukatsu, 1952). A rapid increase in nematode numbers takes place at later tillering (Goto and Fukatsu, 1952) and is associated with the reproductive phase of plant growth (Huang and Huang, 1972).

Nematodes are able to enter spikelets before anthesis, within the boot, and feed ectoparasitically on the ovary, stamens, lodicules and embryo (Dastur, 1936; Huang and Huang, 1972). However, A. besseyi is more abundant on the outer surface of the glumes and enters when these separate at anthesis (Yoshii and Yamamoto, 1950b). As grain filling and maturation proceed, reproduction of the nematode ceases, although the development of J3 to adult continues until the hard dough stage (Huang and Huang, 1972). The population of anabiotic nematodes is predominantly adult females (Huang et al., 1979). These nematodes coil and aggregate in the glume axis. More nematodes occur in filled grain than in sterile spikelets (Yoshii and Yamamoto, 1950b) and infected grain tends to occur more towards the middle of the panicle (Goto and Fukatsu, 1952).

A. besseyi is amphimictic (Huang et al., 1979) and males are usually abundant, however, reproduction can be parthenogenetic. The optimum temperature for oviposition and hatching is 30°C. At 30°C the life cycle is 10±2 days and lengthens significantly at temperatures <20°C (Huang et al., 1972). No development occurs below 13°C (Sudakova, 1968).

A. besseyi aggregate in the glume axis of maturing grain and slowly desiccate as kernel moisture is lost. They become anabiotic and are able to survive for 8 months to 3 years after harvest (Cralley, 1949; Yoshii and Yamamoto, 1950b; Todd, 1952; Todd and Atkins, 1958). Survival is enhanced by aggregation and a slow rate of drying (Huang and Huang, 1974), but the number (Yoshii and Yamamoto, 1950b; Sivakumar, 1987a) and infectivity (Cralley and French, 1952) of nematodes is reduced as seed age increases. It is ironic that good seed storage conditions probably prolong nematode survival.

A. besseyi is not thought to survive long periods in soil between crops (Cralley and French, 1952; Yamada et al., 1953) although anabiotic nematodes may survive on rice husks and plant debris. Sivakumar (1987b) found A. besseyi reproducing on Curvularia and Fusarium in straw after harvest.

The principal dispersal method for A. besseyi is seed. It can be transmitted in flood water in lowland rice (Tamura and Kegasawa, 1958) but the survival of nematodes in water decreases as temperature increases from 20 to 30°C (Tamura and Kegasawa, 1958). High seeding rates in infected seed beds facilitates local dispersal (Kobayashi and Sugiyama, 1977).

Strawberry

A. besseyi is a foliar pest of strawberry and may be found between leaves in buds. The nematode has rapid life cycles (2-3 weeks) and thrive in moist conditions which enable them to move over plant surfaces in water films (Brown et al., 1993).

Natural enemies

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Natural enemy	Type	Life stages	Specificity	References	Biological control in	Biological control on
Arachnula impatiens	Pathogen
Vampyrella vorax	Pathogen

Seedborne Aspects

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Incidence

Surveys have shown large numbers of seed lots to be infected with, and high incidences of infection by, A. besseyi throughout the main rice producing areas of the world. In Tanzania, A. besseyi was reported in 12.8% of rice seed lots with infection levels ranging from 2 to 82% within lots (Taylor et al., 1972). About 70% of the samples of rice grains from 32 plantations in central west Brazil were infected. The numbers of nematodes ranged from 10 to >140 nematodes/100 grains (Huang et al., 1977). A. besseyi was recovered from 5.5% of 474 seed samples obtained from rice seed warehouses in Louisiana, USA (McGawley et al., 1984). In Malaysia, 80% of seed lots from four major rice growing areas showed the presence of A. besseyi (Rahmin, 1988). A. besseyi has also been found in the seeds of Stylosanthes hamata (Gokte et al, 1992), Fraxinus americana (Gokte et al., 1989) and Panicum maximum (Merny et al., 1985).

The process of infection of the seeds begins after sowing, when anabiotic A. besseyi rapidly become active and are attracted to meristematic areas of the developing seedling (Nandakumar et al., 1975). During early growth, it is found in low numbers within the innermost leaf sheath, feeding ectoparasitically around the apical meristem (Yoshii and Yamamoto, 1950b; Goto and Fukatsu, 1952; Todd and Atkins, 1958). The main stem is frequently more infected than subsequent tillers (Goto and Fukatsu, 1952). A rapid increase in nematode numbers takes place at later tillering (Goto and Fukatsu, 1952) and is associated with the reproductive phase of plant growth (Huang and Huang, 1972).

Nematodes are able to enter spikelets before anthesis, within the boot, and feed ectoparasitically on the ovary, stamens, lodicules and embryo (Dastur, 1936; Huang and Huang, 1972). However, A. besseyi is more abundant on the outer surface of the glumes and enters when these separate at anthesis (Yoshii and Yamamoto, 1950b). As grain filling and maturation proceed, reproduction of the nematode ceases, although the development of J3 to adult continues until the hard dough stage (Huang and Huang, 1972). The population of anabiotic nematodes is predominantly adult females (Huang et al., 1979). These nematodes coil and aggregate in the glume axis. More nematodes occur in filled grain than in sterile spikelets (Yoshii and Yamamoto, 1950b) and infected grain tends to occur more towards the middle of the panicle (Goto and Fukatsu, 1952).

A. besseyi aggregate in the glume axis of maturing grain and slowly desiccate as kernel moisture is lost. They become anabiotic and are able to survive for 8 months to 3 years after harvest (Cralley, 1949; Yoshii and Yamamoto, 1950b; Todd, 1952; Todd and Atkins, 1958). Survival is enhanced by aggregation and a slow rate of drying (Huang and Huang, 1974), but the number (Yoshii and Yamamoto, 1950b; Sivakumar, 1987a) and infectivity (Cralley and French, 1952) of nematodes is reduced as seed age increases. It is ironic that good seed storage conditions probably prolong nematode survival.

Togashi and Hoshino (2001) investigated relationships between the number of A. besseyi per seed, size of seed harbouring nematodes, and nematode mortality on 18 panicles collected from 12 paddy fields (in Japan) showing three different levels of white tip disease. The panicles were investigated after storage for 30-82 days at 4°C. There was no nematode mortality within seeds during storage. Mean nematode number per seed increased and mean degree of seed swelling decreased as the paddy field infestation of white tip disease increased. For paddy field means, there was a significant negative correlation between the mean nematode number per seed and the mean degree of rice seed swelling. Intriguingly, for individual seeds, the mean degree of swelling increased up to that typical of well-developed seeds with increasing nematode number per seed. Nematode mortality occurred in an inversely density-dependent fashion within individual seeds. The nematode exhibited a clumped distribution among seeds in each paddy field. Such ecological features of the nematode might contribute to its persistence in rice plant populations.

Effect on Seed Quality

Seeds infected with A. besseyi are capable of germination, but hot water treatment to control the nematode reduced germination of some varieties (Rahim, 1988). According to Tsay et al. (1998), 16% of A. besseyi survived under desiccation at 70°C for 12 h, and the germination of rice seed decreased to 44%. At 60°C, 40% of A. besseyi survived and there was no effect on rice seed germination.

Pathogen Transmission

Seed

The principal dispersal method for A. besseyi is seed. Nandakumar et al. (1975) showed that the nematodes were activated from their dormant state by soaking infested rice seeds in water at 28°C for 12 to 15 hours. They then feed on the tender primordium of the sprouting seeds but later migrate to the leaf as it unfurls. Symptoms of white-tip disease appear on leaves 16 days after germination. A. besseyi adapts to seed transmission through its capacity to remain quiescent upon dehydration and reactivate with rehydration. In northern Italy, where conditions were unfavorable for the overwinter survival of A. besseyi, phytosanitary measures such as the exclusion of infested seeds were effective in reducing the spread of the nematode in paddy crops (Bergamo et al., 2000).

The number of studies undertaken on crop losses due to the nematode is very limited, but an inverse correlation between the number of nematodes in seeds and crop performance has been convincingly demonstrated (Huang, 1983). Generally, population densities per seed number or weight are counted. Fukano (1962) determined an economic damage threshold density (300 live nematodes/100 seed). It has been shown that high seeding rates in infected seed beds facilitates local dispersal of the pathogen (Kobayashi and Sugiyama, 1977).

Other sources

A. besseyi can be transmitted in flood water in lowland rice (Tamura and Kegasawa, 1958) but the survival of nematodes in water decreases as temperature increases from 20 to 30°C (Tamura and Kegasawa, 1958). The nematode also lives in rice stubble left in the field after harvest which enables transfer of the inoculum from season to season (Sivakumar, 1987).

Seed Treatment

There are numerous variations of methods for the hot-water treatment of rice seed (Cralley, 1949, 1952; Yoshii and Yamamoto, 1950c, 1951; Todd and Atkins, 1958; Borokova, 1967). The most effective control of A. besseyi requires seed to be pre-soaked in cold water for 18-24 hours and immersed in water at 51-53°C for 15 minutes. Higher temperatures (55-61°C for 10-15 minutes) are required if seed is not pre-soaked. The temperature and duration of treatment must be closely monitored and, after treatment, the seed must be dried at 30-35°C, or sun-dried if stored, but otherwise can be sown directly in the field. Garcia et al. (2000) eradicated A. bessseyi by heat treatments, wet (60°C/10 minutes and 57°C/15 minutes) and dry (90°C/6 and 12 h and 95°C/6 and 12 h). Hoshino and Togashi (2000) showed that nematicides alone caused little nematode mortality within seeds. Most mortality occurred while seeds were being air-dried. Mortality caused by air-drying alone was 1.7 times greater than that caused by soaking seeds in water for 24 hours.

For quarantine purposes at the International Rice Research Institute (IRRI), seed is soaked in cold water for 3 hours followed by hot water at 55°C for 15 minutes. Tenente and Manso (1994) studied rice seeds infested with A. besseyi either immersed in water at 52, 54 and 57°C for 10 and 15 minutes, with and without shaking, or treated with water solutions at 32 and 33 ml/100 ml of thiabendazole (40%). Thiabendazole decreased the nematode population. Heat treatment with shaking at all temperatures and the 57°C/10 minute treatment without shaking also allowed eradication. Only the heat treatment at 52°C for 15 minutes with shaking gave eradication of A. besseyi without affecting germination, and is thus considered as the most appropriate control method. Giudici et al. (2004) found that seed immersion for 10-15 minutes in water at 55-61°C eradicated seedborne A. besseyi without reducing germination.

Various chemical seed treatments have been used for the control of A. besseyi including demeton, malathion and fensulfothion. Lee et al. (1972) reported effective control by treating water or by root dipping with diazinon and nemagon. A. besseyi control with carbosulfore sprays has been reported (Rao et al., 1986), but pre-harvest chemical treatment alone is only partially effective (Aleksandrova, 1981). The timing of application is important. 

Ultrasound and gamma irradiation have also been investigated for their potential in eradicating A. besseyi from seeds (Aleksandrova 1985; Nagy, 1987).

Seed treatment with dipotassium hydrogen phosphate, coupled with a foliar spray of salicylic acid effectively managed A. besseyi in the field in Karnataka, India, and increased the yield by 3.59% (Ramakrishnam and Raiendran, 2003a). Seed treatment with Pseudomonas flourescens, followed by foliar applications of P. flourescens at 45, 55 and 65 days after sowing, reduced chaffiness in rice (caused by A. besseyi) by 56.5% (Ramakrishnam and Raiendran, 2003b). Mohanty et al. (2004) recommend soaking rice seeds in 0.075% cartap hydrochloride for 12 hours, which resulted in 82.02% mortality of seedborne A. besseyi and a significant increase in grain yield.

Biocontrols were trailed in Ghana by Osei and Sackey-Asante (2006). Seed dressing treatment with ground pepper (Capsicum frutescens cv. Legon 18), ground neem seeds (Azadirachta indica) and wood ash, all at 15 g/kg, reduced seedborne populations of A. besseyi by 83.5%, 82.4% and 71.7%, respectively. However, the treatments also inhibited seed germination by 9.6-24.4%, with ground pepper resulting in the greatest reduction.

Seed Health Tests

Manual dehulling (Huang, 1983; Prot and Gergon, 1994)

- Soak seeds in water for 24 hours.
- Dehull seeds with a scalpel and needle.
- Transfer contents (kernels, hulls and water) to a Baermann funnel or a sieve.
- Recover and count nematodes after 48-72 hours.

Modified Baermann or Seive method (Gergon and Mew, 1991; Prot and Gergon, 1994)

- Place seeds over a 10 cm, 40 x 40 mesh steel wire dish in a funnel (12 cm in diameter) and filled with 250 ml water.
- Let it stand for at least 48 hours.
- After incubation, draw ca 20 ml of water into a test tube through a rubber tube attached to the bottom of the funnel.
- Allow the collected water to stand for 1 hour.
- Pipette out excess water and leave 10-15 ml in the test tube.
- Examine the remaining water for nematodes.

Notes on Methods

A mass extraction method has been described (Hoshino and Togashi, 2002). Common methods of nematode extraction applied to Panicum maximum seeds infested with A. besseyi have been compared by Bueno et al. (2002).

Pathway Vectors

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Vector	Notes	Long Distance	Local	References
Containers and packaging - wood		Yes
Land vehicles		Yes
Mail	Rice seeds	Yes
Plants or parts of plants	Grasses in paddy fields	Yes
Soil, sand and gravel		Yes

Plant Trade

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Plant parts liable to carry the pest in trade/transport	Pest stages	Borne internally	Borne externally	Visibility of pest or symptoms
Bulbs/Tubers/Corms/Rhizomes	adults; eggs; juveniles	Yes	Yes	Pest or symptoms not visible to the naked eye but usually visible under light microscope
Flowers/Inflorescences/Cones/Calyx	adults; eggs; juveniles	Yes	Yes	Pest or symptoms not visible to the naked eye but usually visible under light microscope
Leaves	adults; eggs; juveniles	Yes	Yes	Pest or symptoms not visible to the naked eye but usually visible under light microscope
Stems (above ground)/Shoots/Trunks/Branches	adults; eggs; juveniles	Yes	Yes	Pest or symptoms not visible to the naked eye but usually visible under light microscope
True seeds (inc. grain)	adults; eggs; juveniles	Yes	Yes	Pest or symptoms not visible to the naked eye but usually visible under light microscope

Plant parts not known to carry the pest in trade/transport
Bark
Fruits (inc. pods)
Growing medium accompanying plants
Roots
Seedlings/Micropropagated plants
Wood

Impact

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Rice

A. besseyi is widely distributed because of its dissemination in seed, but its importance varies between regions, countries and localities. Within a locality, the incidence and severity of the disease can change from year to year, and is strongly influenced by cultural practices and local rice types. Infection and damage are generally greater in lowland and deep water systems than in upland environments. However, losses of up to 50% have been reported in upland rice in Brazil (da Silva, 1992).

Damage in a susceptible cultivar largely depends on the percentage of infested seed sown and the number of A. besseyi/infested seed. With few exceptions, the former has rarely been determined despite its importance in governing the number of infection loci in a field. Generally, population densities/seed number or weight are counted. Fukano (1962) determined an economic damage threshold density (300 live nematodes/100 seed).

In the 1950s typical figures for susceptible cultivars in the USA were 17.5, 4.9 and 6.6% in different years (Atkins and Todd, 1959) and 10-30% in Japan (Yamada and Shiomi, 1950; Yoshii and Yamamoto, 1950a; Yoshii, 1951). Tsay et al. (1998) reported yield losses of 44.9, 34.7 and 24.2% when rice plant infestation rates were 57, 34 and 18%, respectively. A. besseyi has been controlled in the USA by seed treatment and the use of resistant cultivars and is no longer a pest (Hollis and Keoboonrueng, 1984). A. besseyi also disappeared from Japan but has re-occurred, the economic value of infected discoloured grain being reduced if infection exceeds 0.7% (Inagaki, 1985).

A. besseyi damage has been reported from deep water rice in Bangladesh. More than 50% of fields were infected and the panicle weight of heavily infected plants (650 nematodes/100 seed) was a third that of less infected plants (112 nematodes/100 seed) (Rahman and McGeachie, 1982; Rahman and Taylor, 1983). In contrast, local cultivars in Thailand appear to be tolerant of A. besseyi and no symptoms have been observed despite widespread infection (Buangsuwon et al., 1971).

Economic loss in the Philippines has not been reported, but infection varies according to year, season and cultivar (Madamba et al., 1974). Levels of infested seed are generally low (4.7-7% over 5 years) (Madamba et al., 1981), and severe damage is unlikely as high numbers of A. besseyi (210-5300/100 seed) are not always associated with a high percentage of infested seed.

A. besseyi is thought to be an important pest in India. Rao (1976) reported severe symptoms in the field, but accurate yield loss assessment is lacking. Muthukrishnan et al. (1974) observed that plants sometimes recover after early severe damage and computed losses of 0.2-10%. Infestation levels in Sri Lanka are not considered important (Lamberti and Robini, 1980).

Strawberry

A. besseyi is an important pest of strawberry in the USA, south of Arkansas and Virginia (Brown et al., 1993).

Diagnosis

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A diagnostic protocol for A. besseyi is given in OEPP (2004). The diagnosis of A. besseyi from abnormal rice grains in China is detailed in Lin et al. (2005).

A. Besseyi was identified in forage grasses in Brazil by observing morphological features under the light and scanning electron microscopes in Favoreto et al. (2011).

Detection and Inspection

Top of page Different sampling methods are used for A. besseyi, depending on the stage of crop growth. During early growth and tillering, A. besseyi is found in the base of the culm and between leaf sheaths. For immediate inspection, plant tissue can be teased in water to release the nematodes. This method is suitable for tissue of any potential host species. Plant tissue can be stained before examination; this is particularly useful for detecting low numbers. Alternatively, A. besseyi can be extracted from chopped tillers placed on a sieve, or directly in water. During the reproductive phase in rice, A. besseyi is recovered from spikelets and grain by soaking a known number in water for 24-48 h at 25-30°C. Quantitative extraction from rice requires that the glumes are separated from the kernel but remain in the extract. The percentage of infested seed is a useful parameter, but extracting from individual seeds is time consuming. Better recovery is achieved from hulled grain, but extraction from unhulled grain is sufficient for detection of A. besseyi (e.g. for quarantine) from a large seed sample.

Similarities to Other Species/Conditions

Top of page A. besseyi could be confused with Ditylenchus angustus.

Prevention and Control

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Preventing the dispersal of A. besseyi in rice requires the elimination of nematodes from seed; hot water or chemical seed treatments are commonly used. Resistant cultivars and cultural methods have been used to reduce infection below damage thresholds, and tolerant cultivars avoid yield loss without nematode control. Stubble burning prevents transmission of A. besseyi in straw and chaff but would have to be used in conjunction with other control measures.

Hot water treatment

There are numerous variations of methods for the hot water treatment of rice seed (Cralley, 1949, 1952; Yoshii and Yamamoto, 1950c, 1951; Todd and Atkins, 1958; Borokova, 1967). The most effective control requires seed to be pre-soaked in cold water for 18-24 hours, then immersed in water at 51-53°C for 15 minutes. Higher temperatures (55-61°C for 10-15 min) are required if seed is not pre-soaked. The temperature and duration of treatment must be closely monitored and, after treatment, the seed must be dried at 30-35°C, or sun dried if stored, but otherwise can be sown directly in the field. For quarantine purposes at the International Rice Research Institute, seed is soaked in cold water for 3 hours followed by hot water at 55°C for 15 minutes.


Chemical Control

Various chemical seed treatments have been used for the control of A. besseyi including demeton, malathion and fensulfothion. Lee et al. (1972) reported effective control by treating water or by root dipping with diazinon and nemagon. A. besseyi control with carbosulfore sprays has been reported (Rao et al., 1986), but pre-harvest chemical treatment alone is only partially effective (Aleksandrova, 1981). The timing of application is important. 

Host-Plant Resistance

Resistance to A. besseyi is widespread. In the USA, A. besseyi has been controlled principally through the use of resistant cultivars. Resistance to A. besseyi has subsequently been reported from Japan (Nishizawa, 1953b); Yamada et al., 1953; Goto and Fukatsu, 1956), Korea (Park and Lee, 1976), India (Rao et al., 1986), Brazil (Oliveira and Ribeiro, 1980; da Silveira et al., 1982), USSR (Popova et al., 1980) and Italy (Orsenigo, 1954). Resistance to A. besseyi is thought to be genetically controlled and carried by the Japanese cultivar Asi-Hi (Nishizawa, 1953).

Symptomless but susceptible (i.e. tolerant) cultivars (Nishizawa, 1953; Goto and Fukatsu, 1956) are common. Symptom expression in the field is particularly variable (Atkins and Todd, 1959) and variations between plants of a cultivar also occur (Orsenigo, 1954). In Thailand, all local cultivars are considered tolerant of A. besseyi (Buangsuwon et al., 1971).


Cultural Control

Irrigating seed beds (Yamada et al., 1953) or direct seeding into water (Cralley, 1956) reduces infection by A. besseyi. In these conditions, nematodes emerge and lose vigour before seed germination. High seedling rates in the seed bed (Kobayashi and Sugiyama, 1977) and high numbers of seedlings/hill (Yamada et al., 1953) tend to increase infection by increasing the number of infection loci in the field. Such problems are thought to be responsible for the re-occurrence of A. besseyi in Japan (Inagaki, 1985). In the USA (Cralley, 1949) and Japan (Yoshi and Yamamoto, 1951; Yamada et al., 1953), early planting, presumably in cooler conditions, reduced or eliminated A. besseyi infection.

Summary of Control Measures

- Hot water treatment of seed. Probably the most effective and cheapest control measure

- Resistant or tolerant cultivars

- Early planting if rice season is preceded by a cooler period

- Low seed bed planting densities

In strawberry, control is possible by hot water treatment of planting stocks, or regeneration of plants from dormant excised axillary buds. Host weed control and planting healthy runners in clean soil is the basis for ensuring freedom from problems.

References

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