Rev 01/09/2024
Tylenchida Aphelenchina Aphelenchoidea Aphelenchoididae Aphelenchoidinae
Aphelenchoides besseyi Christie, 1942
Dorsal esophageal gland opens into lumen of esophagus in metacorpus, just anterior to the pump chamber.
Subventral esophageal glands open into lumen of esophagus in metacorpus, posterior to the pump chamber.
From Franklin and Siddiqi, 1972
The nematode has a slender stylet with small, distinct knobs.
Females: 0.66-0.75 mm long and slender.
Males: are as numerous as females.
However, note that the classification based on morphological characters is not well supported by an integrated approach including both molecular and phylogenetic analysis. Ratherr, A. besseyi sensu lato appears to be a species complex with several cryptic species that are not well delimited by morphology (Subbotin et al., 2021).
Species delimitation among A. besseyi sensu lato. populations is mportant for phytosanitary purposes and for the selection of crop cultivars resistant to these nematodes. For example, A. besseyi sensu stricto parasitizes only strawberry (Olivieria et al., 2019). whereas A. besseyi sensu lato has a much wider host range. Conseqquently, Subbotin et al (2021) proposed the reinstatemnt of A. oryzae, which was originally described by Yokoo (1948) as the causal agent of white tip of rice but later synonymized with A, besseyi by Allen (1952).. Subbotin et al., (2021) also proposed A. pseudobesseyi and A. pseudogoodeyi for variants of A. besseyi sensu lato that infect cotton and soybeans (Scheck, 2021)..
Reported median body size for this species (Length mm; width micrometers; weight micrograms) - Click:
Aphelenchoides besseyi is widely distributed and occurs in most rice growing areas (Ou, 1985). The known distribution of A. besseyi on rice includes; Australia, Ceylon, Comoro Islands, Cuba, El Salvador, Hungary, India, Indonesia, Italy, Japan, Madagascar, Mexico, Pakistan, Philippines, Taiwan, Thailand, former USSR, and in most countries of central and West Africa (Ou, 1972; Franklin and Siddiqi, 1972). Aphelenchoides besseyi has also been reported as the causal agent of false angular leafspot of beans in Costa Rica (Barrantes et al., 2006; Salas and Vargas, 1984).
Aphelenchoides besseyi was first found in Japan in 1915. In the USA the disease was first noticed in the southern United States in 1935 but was attributed to a nutrient deficiency (Ou, 1972).
Aphelenchoides besseyi has been reported in many Southern U.S. states that produce rice.
White tip disease caused by Aphelenchoides besseyi, is an economic problem in many countries. Recent detections of this nematode in California have sparked the concern of California's rice industry.
From 1959 to 1996, A. besseyi was detected only twice in California by CDFA Nematologists: once in 1959, in a quarantine strawberry sample originating in Oregon, and the second time in 1963 in a fungal culture collected in Butte county. Attempts to find the nematode again from the same field were unsuccessful. A survey of the state by CDFA for the presence or absence of A. besseyi in California paddy rice was initiated in 1997. It was intended that this survey would provide a sound basis for certifying California paddy rice free of A. besseyi, and thereby, eliminate the requirement by the government of Turkey for methyl bromide treatments of export shipments. Sampling was designed to detect the presence of the nematode at the county or rice growing region level and 170 samples were collected.
During the 1997 survey, one confirmed and three suspected findings of A. besseyi resulted in four samples collected from two counties. These samples tested negative when examined a second time. However, the government of Turkey required that batches of rice intended for shipment to that country should be sampled and certified to be free of the nematode. Between 1998 and 2001, A. besseyi has been found in three such export loads. Those shipments were rejected, resulting in millions of dollars in losses.
A-rated pest in California Nematode Pest Rating System.
At the state level there are no formal quarantine regulations implemented against A. besseyi. There are no prohibitions for the import of rice seed into California from other states within the US. At the federal level, federal regulations have prohibited the import of seed and paddy rice into the US since November 23, 1933. Milled rice can enter California.
Paddy rice exported to Turkey requires certification for freedom from A. besseyi according to the Turkish Government pest control requirements. In compliance with the Turkish government, certification is given on a per shipment basis; each shipment has to be sampled and found free of A. besseyi before it can be certified and shipped to Turkey.
False angular leafspot of beans is considered an important emergent disease of beans in Costa Rica. In many areas of Costa Rica beans are grown in rotation with rice (Barrantes et al., 2006).
Although A. besseyi is widely distributed in Brazil, economic losses have rarely been reported. However, the frequent association of A. besseyi with forage grass seeds in Brazil represents a phytosanitary barrier for export to many countries. In addition, other morphologically similar but non-pathogenic Aphelenchoides species are also associated with forage grass seeds, leading to difficulty in attaining a robust diagnosis (de Jesus et al., 2016).
Mainly ectoparasitic on buds.
Feeds at leaf tips and growing points in rice.
May feed endoparasitically in leaf mesophyll.
Rice is the most important host world wide.
On strawberry A. besseyi is the causal agent of 'summer dwarf' or 'crimp' in the United States and Australia.
Other host plants include; onion, garlic, sweet corn, sweet potato, soybean, yan chinese cabbage, sugar cane, horseradish, lettuce, millet, many grasses, orchids, wishbone, chrysanthemum, marigold, mexican sunflower, african violets, rubber plant, Hibiscus brachenridgii, and hydrangea (Franklin and Siddiqi, 1972). Aphelenchoides besseyi has also been reported as the causal agent of false angular leafspot of beans in Costa Rica (Barrantes et al., 2006; Salas and Vargas, 1984).
The following are listed as hosts in various literature sources:
Ecophysiological Parameters:
Aphelenchoides besseyi is a seed-borne ectoparasite rice. Nematodes in dry rice seeds are iin anhydrobiosis and are activated by rehydration. Hoshino and Togashi (2020) demonstrated that Survival rate in dry rice seeds was lower at 20 and 25 C than at −5 to 10 C after storage for 190 days. No nematodes survived a storage period of 1313 days at 20 and 25 C.
However, when rice seeds were held at 5 C, the survival rate of nematodes decreased during a period of 7315 days. In storage at −30 C, nematode survival was not affected during a period of 6485 days. After storage at −30 C for 6546 days, 96% of rice seeds germinated; 60-96% of seeds germinated after storage at 5 C for 6610-7373 days. Therefore, Hoshino and Togashi (2020) postulated that low temperature storage of rice seeds (between %C and -30C) would not signicicantly disrupt the pathogenicituy of A, besseyi on rice.
As a non-obligate phytoparasite, A. besseyi can survive in the absence of the host plant as a fungal-feeder in the soil; under extreme dehydration it can survive for a long time in an anhydrobiotic state inside plant debris or seeds. In the anhydrobiotic in dry tissues, under hulls of rice grains; the nematode is viable after 3 years.
This nematode is not thought to survive long periods in soil between crops (Cralley and French, 1952).
This nematode is mainly ectoparasitic feeding on young tissue. At the end of the growing season many nematodes are in a state of cryptobiosis under the hulls of seed (Taylor, 1969).
Seed infected with Aphelenchoides besseyi is planted and the nematodes become active and are attracted to the meristematic areas. During early growth, A. besseyi is found in low numbers within the folded leaf sheath, feeding ectoparasitically around the apical meristem (Todd and Atkins, 1958). A rapid increase in nematode numbers takes place at late tillering and is associated with the reproductive phase of the plant. Nematodes are able to enter the spikelets before anthesis and feed ectoparasitically on the plants reproductive structures. As grain filling and maturation proceed, reproduction of the nematode ceases, although the development of third stage juveniles to adult continues until the hard dough stage.
The population of nematodes is predominately adult female which are normally amphimictic, although parthenogenetic reproduction has been reported (Sudakova and Stoyakov, 1967).
These nematodes coil and aggregate in the glume axis.
The optimum temperature for oviposition and hatch is 30C.
At 30 C the life cycle is approximately 8-12 days and no development occurs below 13C (Sudakova, 1968).
Aphelenchoides besseyi slowly desiccate as kernel moisture is lost. They become anhydrobiotic dormant, and are able to survive in a quiescent state on infested seed for long periods of time, from 8 months to 3 years (Cralley, 1949; Yoshi and Yamamoto, 1950). Survival is enhanced by aggregation and a slow rate of drying (Huang and Huang, 1974), but the infectivity of the nematode is reduced as seed age increases (Cralley and French, 1952).
In the rice seed-bed, emergence of severely infected seedlings is delayed and germination is low.
The most conspicuous symptoms occur at the tillering stage (Taylor, 1969).
Yields of rice may be reduced 50%.
Experiments have shown that different varieties of rice are affected to different degrees of infestation. Yields have been reported to be reduced by 17-54% in susceptible varieties and 24% in resistant (Atkins and Todd, 1959).
Aphelenchoides besseyi is reported to be the causal agent of the green stem and retention foliar syndrome (GSFR) in soybean (Meyer et al., 2017) and infecting shoots of cotton plants (Favoreto et al., 2018; Calandrelli et al., 2023)
Host Plant Resistance, Non-hosts and Crop Rotation alternatives:
Hot water treatment of seed can be used to destroy this nematode infecting the seeds (Atkins and Todd, 1959). Pinherio et al. (1997), found that thermal wet treatment was the most effective. Aphelenchoides besseyi was not recovered from rice seeds which received hot water treatment at 55-60C for 15 minutes, but was observed in seed subjected to hot water treatment at 50C (Gergon and Prot, 1993). Lower temperatures (52-53C for 15 minutes) for hot water treatment have been reported (Crawley, 1959; Tenente et al., 1994). At temperatures reported for controlling A. besseyi, no effect on seed germination or viability was reported if seed was planted quickly. Chemical treatments of seed have also been reported as being effective in controlling A. besseyi. Benomyl applied as a seed treatment reduced average numbers of nematodes (Gergon and Prot, 1993; Templeton et al., 1971). Thiabendazole has also been reported as an effective seed treatment (Tenente and Manso, 1994). Carbofuran (Tenente and Manso, 1994; Todd and Atkins, 1959) and aluminum phosphate fumigation (Tenente et al., 1994), are not effective chemical seed treatments for A. besseyi.
Allen, M.W. 1952. Taxonomic status of the bud and leaf nematodes related to Aphelenchoides fragariae (Ritzema Bos, 1891). Proc. Helminth. Society of Washashington. 19:108-120.
Barrantes, W., Araya, C.M., Esquivel, A. 2006. Falsa mancha angular del frijol: una enfermedad que avanza en Costa RicaManejo Integrado de Plagas y Agroecologia (Costa Rica) No. 78, 2006
Bridge, J., M. Luc and R.A. Plowright. 1990. "Nematode parasites of rice." Pp. 69-108, M. Luc et al. (eds.), Plant parasitic nematodes in subtropical and tropical agriculture. Wallingford, UK: CAB International.
Calandrelli, A., Silva, M.T.R., Miamoto, A., Rinaldi, L.K. , Favoreto, L., Meyer, M.C., Machado, A.C.Z., Silva, S.A., Dias-Arieira, C.R. 2023. Host-parasite relationship between Aphelenchoides besseyi and soybean, Nematology 25:33-44.
Cheng, X., Xiang, Y., Xie, H., Xu, C.-L., Xie, T.-F., Zhang, C. & Li, Y. (2013). Molecular characterization and functions of fatty acid and retinoid binding protein gene (Ab-far-1) in Aphelenchoides besseyi. PLoS ONE 8, e66011.
Cralley, E.M. 1949. "White tip of rice." Phytopathology. 39:5.
Cralley, E.M., and R.G. French. 1952. "Studies on the control of white tip of rice." Phytopathology. 42:6.
de Jesus, D.S., Gon alves Oliveira, C.M., Roberts, D., Blok, V., Neilson, R., Prior, T., Balbino, H.M., MacKenzie, K.M. D Arc de Lima Oliveira, R. 2016. Morphological and molecular characterisation of Aphelenchoides besseyi and A. fujianensis (Nematoda: Aphelenchoididae) from rice and forage grass seeds in Brazil. Nematology 18:337 356.
Favoreto, L., Faleiro, O., Freitas, M.A., Brauwers, L.R., Galbieri, R., Homiak, J.A., Lopes-Caitar, V.R., Marcelino-Guimar�es, FC. & Meyer, M.C. 2018. First report of Aphelenchoides besseyi infecting the aerial part of cotton plants in Brazil. Plant Disease 102, 2662. DOI: 10.1094/PDIS-02-18- 0334-PDN
Franklin, M.T. and M.R. Siddiqi. 1972. Aphelenchoides besseyi. Commonwealth Institute of Helminthology Descriptions of Plant-parasitic Nematodes Set 1, No. 4.
Gergon, E.B. and J.C. Prot. 1993. "Effect of benomyl and carbofuran on Aphelenchoides besseyi on rice." Fundamentals of Applied Nematology. 6:563-566.
Gokte, N. and V. K. Mathur. 1993. Treatment schedule for denematization of seeds of Setaria italica and Panicum miliaceum infested with Aphelenchoides besseyi. Nematologica 39:274-276.
Goodey, J. B., M. T. Franklin, and D. J. Hooper. 1965. T. Goodey's: The Nematode Parasites of Plants Catalogued Under Their Hosts. Commonwealth Agricultural Bureaux, Farnham Royal, Bucks, England. Third Edition
Hoshino, S. and Togashi, K. 2020. Effects of temperatures on survival of Aphelenchoides besseyi in prolonged storage of rice grains. Nematology 22:1169-1177.
Huelma, C.C., J.C. Prot, S.D. Merca, and T.W. Mew. 1994. "Aphelenchoides besseyi in irrigated upland and lowland rice during dry and wet season." International Rice Research Notes. 19 (3):30.
Ibrahim, S. K., R. N. Perry and D. J. Hooper. 1994. Use of esterase and protein patterns to differentiate two new species of Aphelenchoides on rice from other species of Aphelenchoides and from
Meyer, M.C., Favoreto, L., Klepker, D. & Marcelino-Guimar�es, F.C. 2017) Soybean green stem and foliar retention syndrome caused by Aphelenchoides besseyi. Tropical Plant Pathology 42, 403-409. DOI: 10.1007/s40858-017-0167-z
Ou, S.H. 1985. Rice Diseases (2nd edition). New England: Commonwealth Mycological Institute.
Pinheiro, F.P., R.P. Vianello, F.S. Ebeidalla, and R.C.V. Tenente. 1997. "Thermal seed treatments to eradicate Aphelenchoides from Brachiaria dictyoneura." Nematologia Brasilerira. 21 (1): 92-97.
Salas, LA; Vargas, E. 1984. El nematodo foliar Aphelenchoides besseyi Christie (Nematode: Aphelenchoididae) como causante de la falsa mancha angular del frijol de Costa Rica. AgronomÃÂa Costarricense 8:65-68
Sanwal, K.C. 1961. A key to the species of the nematode genus Aphelenchoides Fischer, 1894. Canadian Journal of Zoology. 39:143-148.
Scheck, H.J. 2021. California Pest Rating Proposal for Aphelenchoides besseyi Christie, 1942 Strawberry summer crimp nematode. CDFA Sacramento, California, USA.
Subbotin, S.A., Oliveira, C.J., Alvarez-Ortega, S., Desaeger, J., Crow, W., Overstreet, C., Leany, R., Vau, S. and Inserra, R.H. 2021. The taxonomic status of Aphelenchoides besseyi Christie, 1942 (Nematoda: Aphelenchoididae) populations from the Southeastern USA, and description of Aphelenchoides pseudobesseyi sp. n. Nematology 23: 381-413
Taylor, A.L. 1969. "Nematode parasites of rice." Pp. 264-268, J.E. Peachey (ed.), Nematodes of Tropical Crops. Albans, Herts, England: Commonwealth Bureau of Helminthology.
Tamura, I., and K. Kegasawa. 1958. "Studies on the ecology of the rice nematode, Aphelenchoides besseyi Christie. II. On the parasitic ability of rice nematodes and their movement into hills." Japanese Journal of Ecology. 8:37-42.
Templeton, G.E., T.H. Johnston and J.T. Daniel. 1971. Benomyl controls rice white tip disease." Phytopathology. 61:1522-1523.
Tenente, R.C.V., M.M.V.S. Wetzel, E. S. Manso, and A.S. Marques. 1994. "Survival of Aphelenchoides besseyi in infested rice seeds stored under controlled conditions." Nematologia Brasiliera. 18 (1): 85-92.
Tenente, R.C., and E.S. Manso. 1994. "Chemical and heat treatments of rice seeds infested with Aphelenchoides besseyi." Nematologia Brasileira. 18 (1): 28-34.
Yokoo, T. 1948. [Aphelenchoides oryzae Yokoo n. sp. a nematode parasite to rice plant.] Annals of the Phytopathological Society of Japan 13, 40-43.
Yoshi, H., and S. Yamamoto. 1950. "A rice nematode disease 'Sencha Shingane Byo'. II. Hibernation of Aphelenchoides oryzae." Journal of Faculty of Agriculture, Kyusha University. 9:223-233.