Introduction
Phytophthora megasperma Drechsler (1931)
Phytophthora megasperma was isolated by Drechsler in 1931 from root tissue of hollyhock (Althaea rosea) (Drechsler, 1931). A synonym of P. megasperma is Pythiomorpha miyabeana S. Ito & Nagai (1931) (Cline et al., 2008). Tompkins et al. (1936) broadened Drechsler’s description to include isolates with small oogonia from cauliflower. Waterhouse (1963) separated P. megasperma into P. megasperma var. sojae and P. megasperma var. megasperma based on oogonium size (small versus large, respectively). Kuan and Erwin (1980) questioned the separation of the species based on oogonium size since isolates from different hosts show a continuum of oogonium sizes, and they renamed the species P. megasperma f. sp. glycinea and P. megasperma f. sp. medicaginis for the soybean and alfalfa isolates, respectively. Others also revised the species (Faris et al., 1989). Hansen and Maxwell redescribed P. megasperma in 1991 based on Drechsler’s original description (Hansen and Maxwell, 1991). Isolates of P. megasperma from different hosts were previously given subspecies distinction but are now designated as three separate species, known as P. sojae (soybean), P. medicaginis (alfalfa), and P. trifolii (clover) (Hansen and Maxwell, 1991). The broad host range group of P. megasperma isolates from fruit trees (e.g., cherry, apple) and from the DF1 and soybean race nonclassifiable isolates have recently been separated into the new species P. rosacearum and P. sansomeana, respectively (Cooke et al., 2000; Förster and Coffey, 1993; Hansen et al., 2009). Gallegly and Hong (2008) separate P. megasperma into three subgroups based on the absence (I, II) or presence (III) of hyphal swellings and chlamydospores, oogonial and sporangial size and shape, antheridial characters, and DNA markers. Type II isolates most closely fit the original description (Gallegly and Hong, 2008). See Erwin and Ribeiro (1996) for a thorough explanation of the nomenclature of this complex group. P. megasperma is a group V Phytophthora species (Stamps et al., 1990) (Fig. 1).
Cultural Characteristics
Cultures of P. megasperma are slightly radiate with a medium amount of aerial mycelium (Waterhouse and Waterston, 1966) (Fig. 2). Isolates from hollyhock (Tompkins et al., 1936) grow best at 25°C, and isolates from cauliflower, stock, cabbage, and cineraria are similar. Optimum temperatures for growth for other isolates are Douglas fir (DF1), 27.1°C; DF2, 21.6°C; large-oogonia isolates from alfalfa (A1F2), 22.5°C; rose, 25°C; apple, 22.5°C; grape, 20°C; juniper, 22.5°C; cherry, 25°C; pear, 25°C; poplar, 20°C; and Brassica spp., 20°C.
Reproductive Structures
Asexual Structures
Sporangiophores:
Sporangiophores are simple or sparingly branched, thin, and 2–2.5 µm wide. At the base of the sporangium, sporangiophores can reach widths of up to 5 µm. Sporangiophores usually proliferate inside the empty sporangium (Waterhouse and Waterston, 1966).
Sporangia:
Sporangia are nonpapillate, noncaducous, ovoid, obpyriform, and persistent on the stalk and they proliferate internally (Bush et al., 2006; Drechsler, 1931). Sporangia are 25–45 × 35–60 µm (Fig. 3).
Chlamydospores:
P. megasperma type I and II isolates do not produce chlamydospores. Type III isolates produce chlamydospores.
Hyphae:
Hyphae are 3 µm wide (Waterhouse and Waterston, 1966). Hyphal swellings are rare on solid media but are produced in aqueous cultures of some type III isolates (Gallegly and Hong, 2008). Swellings are rounded or angular and can occur in chains or clusters (Fig. 1).
Sexual Structures
P. megasperma is mostly homothallic, but Barr (1980) and Ho (1986) reported that a large-oospore isolate from alfalfa was heterothallic.
Antheridia:
Antheridia are mainly paragynous, but some amphigynous antheridia form (type I isolates) (Gallegly and Hong, 2008). Antheridia are irregularly spherical or ellipsoid and 10–18 x 14–20 µm.
Oogonia:
Oogonia are produced in both host tissues and axenic cultures. Oogonia are smooth, subspherical, and 42–52 µm in diameter (average 47.4 µm). The oogonia wall is colorless or yellowish, smooth, and up to 1.5 µm thick (Drechsler, 1931). Type II isolates produce the largest oogonia (Gallegly and Hong, 2008).
Oospores:
Oospores are 26–52 µm in diameter (average 41.1 µm) (Drechsler, 1931) (Fig.
4). The oospore wall is smooth and up to 5 µm thick (unstained) or 7 µm thick
(stained).
Host Range and Distribution
The species has a wide host range and can infect many different plant species (Hansen and Maxwell, 1991).
Host |
Common Name |
Disease |
Geographical Distribution |
Actinidia deliciosa |
Kiwifruit |
Root
rot |
|
Aesculus
spp. |
Horse
chestnut |
Root
rot |
|
Althaea rosea |
Hollyhock |
Crown
rot |
|
Asparagus officinalis |
Asparagus |
Soft
rot of spears |
|
Banksia
hookeriana |
Banksia |
Root
rot |
|
Beta vulgaris |
Beet |
Root
rot |
|
Brassica spp. |
Cabbage, cauliflower, Brussels sprouts, turnip, kale, broccoli, rutabaga,
swede, marrow stem, tyfon |
Storage
rot; seedling damping-off; stem rot |
Canada,
Ireland, United Kingdom, United States, Greece, Ireland, Canada |
Capsicum annuum |
Pepper |
Fruit
rot |
|
Carthamus tinctorius |
Safflower |
Root
and stem rot |
|
Chamaecyparis spp. |
Cedar |
Blight |
|
Cheiranthus cheiri |
Wallflower |
Foot
rot |
United
States, |
Cicer arietinum |
Chickpea, garbanzo, gram |
Root
rot |
|
Citrus spp. |
Lemon,
grapefruit, orange |
Root
rot; stem canker |
|
Cucumis sativus |
Cucumber |
Fruit
rot |
|
Dianthus caryophyllus |
Carnation |
Crown
rot |
|
Daucus carota subsp. sativus |
Carrot |
Black
rot; storage or tuber rot |
|
Dryandra polycephala |
Many-headed dryandra |
Root
rot |
|
Fortunella spp. |
Kumquat |
Stem
canker |
|
Hebe spp. |
Hebe |
|
|
Helianthus annuus |
Sunflower |
Stem
rot |
|
Juglans regia |
English
walnut |
Root
and crown rot |
|
Lycopersicon esculentum |
Tomato |
Fruit
rot |
|
Malus
pumila |
Apple |
Root
rot; trunk canker |
|
Matthiola incana |
Stock,
gillyflower |
Root
rot |
|
Medicago sativa |
Alfalfa, l |
Root
rot |
United
States, |
Narcissus
spp. |
Daffodil |
Tuber
rot |
|
Olea europaea |
Common
olive |
Collar
and trunk canker |
|
Oryza sativa |
Rice |
Seedling disease |
|
Persea
americana |
Avocado |
Collar
and trunk canker |
|
Picea abies |
Norway
spruce |
Root
rot |
|
Pinus spp. |
Sugar
pine, pine |
Root
rot; seedling damping-off |
|
Prunus spp. |
Apricot; sweet, sour and Mahaleb cherry; common plum; almond; peach |
Apoplexy; sudden wilt; root and crown rot; trunk canker |
|
Pseudotsuga menziesii |
Douglas
fir |
Seedling rot; chlorosis; stunting |
|
Pyrus malus |
Apple |
Fruit
rot |
|
Rorippa
spp. |
Yellowcress |
Root
rot |
|
|
Rose |
Stem
blight |
|
Rubus idaeus var. idaeus |
Raspberry |
Root
rot |
|
Saccharum officinarum |
Sugar
cane |
Seed-piece rot |
|
Senecio
cruentus |
Cineraria, groundsel |
Root
rot |
|
Solanum spp. |
Eggplant, potato |
Fruit
rot; pink tuber rot |
|
Spinacia oleracea |
Spinach |
Root
rot |
|
Theobroma cacao |
Cacao |
Pod rot |
|
Trifolium repens |
White
clover |
Root
rot |
|
Symptoms
Root Rot of
Brassica spp. (e.g., Cabbage, Cauliflower, Brussels Sprouts, Turnip, Kale,
and Rutabaga):
Infection of the plant starts at the roots and then spreads to the leaves.
Leaves turn progressively purple or reddish from the margins inward, wilt, and
eventually shed. Stems become soft
and flaccid, and the lower end of the taproot blackens and rots.
The cortex sloughs off the taproot and lateral roots, and adventitious roots
often develop above the diseased portions of the taproot.
The disease develops in moist soil at moderate soil temperatures of 20–30°C.
Crown Rot of
Malus pumila (Apple):
Crown rot symptoms include delayed bud break, stunted growth, orange discoloration of the bark, and reduced crop yield (Robertson and Dance, 1971). Crown rot produces cankers that eventually girdle the trunk and cause the bark within to rot completely. Disease symptoms are similar on Prunus spp. (Figs. 5 and 6), except that gum exudes from the cankers. Lesions of various lengths develop on lateral roots, bud break is delayed, and branches die back (Erwin and Ribiero, 1996). The disease develops after prolonged heavy rains.
References
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Phytophthora megasperma.
Bush, E. A., Stromberg, E. L., Hong, C., Richardson, P. A., and Kong, P. 2006.
Illustration of key morphological characteristics of Phytophthora species
identified in
Cline, E. T., Farr,
D. F., and Rossman, A. Y. 2008. A
synopsis
of Phytophthora with accurate scientific names, host range, and
geographic distribution. Plant Health Progress doi:10.1094/PHP-2008-0318-01-RS.
Cooke, D. E. L., Drenth, A.,
Drechsler, C. 1931. A crown rot of hollyhocks caused by
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1996.
Phytophthora Diseases Worldwide. American Phytopathological
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Förster, H., and Coffey, M. D. 1993. Molecular taxonomy of the
Phytophthora megasperma based on mitochondrial and nuclear DNA
polymorphisms. Mycol. Res. 97:1101-1112.
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and DNA Fingerprints. American Phytopathological Society Press,
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Hansen, E. M., and Maxwell, D. P. 1991. Species of the Phytophthora megasperma complex. Mycologia 83:376-381.
Hansen, E. M., Wilcox, W. F., Reeser, P. W., and Sutton, W. 2009. P. rosasearum and P. sansomeana, new species segregated from the Phytophthora megasperma complex. Mycologia 101:129-135.
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Phytophthora
megasperma isolates from soybean and alfalfa.
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14:509-551.
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Institute, Kew,
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Phytophthora de Bary. Mycol. Pap. 92.
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