Figure 1. Morphology of Phytophthora megasperma. Upper row, Ovoid and obpyriform, nonpapillate sporangia; sporangium showing nesting or internal proliferation of a subsequently formed sporangium; sporangium with a new sporangium forming by external proliferation. Lower row, Oogonia, one with a paragynous antheridium and another with an amphigynous antheridium; hyphal swellings; sporangia forming by external proliferation. (Courtesy A. Vaziri; Reproduced from Erwin and Ribeiro, 1996) Click image to see larger view.

 

Figure 2. Culture of Phytophthora megasperma grown on V-8 juice agar. (Courtesy Jean B. Ristaino)

 

Figure 3. Nonpapillate sporangium of Phytophthora megasperma. Bar = 10 µm. (Courtesy Elizabeth A. Bush; Reproduced from Bush, et al., 2006)

 

Figure 4. Oogonium of Phytophthora megasperma with a paragynous antheridium (×1,000). (Courtesy Jean B. Ristaino)

 

Figure 5. Crown rot, caused by Phytophthora megasperma, on an almond tree in Kern County, CA. (Courtesy Greg Browne, USDA-ARS, University of California-Davis)

 

Figure 6. Symptoms of Phytophthora megasperma on almond trees in Kern County, CA. (Courtesy Greg Browne, USDA-ARS, University of California-Davis)

 

 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 

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

United States

Aesculus spp.

Horse chestnut

Root rot

United Kingdom

Althaea rosea

Hollyhock

Crown rot

United States

Asparagus officinalis

Asparagus

Soft rot of spears

New Zealand, United States

Banksia hookeriana

Banksia

Root rot

Australia

Beta vulgaris

Beet

Root rot

United Kingdom

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

Japan

Carthamus tinctorius

Safflower

Root and stem rot

United States

Chamaecyparis spp.

Cedar

Blight

Ireland

Cheiranthus cheiri

Wallflower

Foot rot

United States, Canada

Cicer arietinum

Chickpea, garbanzo, gram

Root rot

Argentina

Citrus spp.

Lemon, grapefruit, orange

Root rot; stem canker

Argentina

Cucumis sativus

Cucumber

Fruit rot

Japan

Dianthus caryophyllus

Carnation

Crown rot

New Zealand

Daucus carota subsp. sativus

Carrot

Black rot; storage or tuber rot

Tasmania, United States, Canada

Dryandra polycephala

Many-headed dryandra

Root rot

Australia

Fortunella spp.

Kumquat

Stem canker

Argentina

Hebe spp.

Hebe

Collar rot

Ireland

Helianthus annuus

Sunflower

Stem rot

Iran

Juglans regia

English walnut

Root and crown rot

United States

Lycopersicon esculentum

Tomato

Fruit rot

United States, Japan

Malus pumila

Apple

Root rot; trunk canker

Italy, New Zealand, United States, Japan

Matthiola incana

Stock, gillyflower

Root rot

United States

Medicago sativa

Alfalfa, lucerne

Root rot

United States, Canada

Narcissus spp.

Daffodil

Tuber rot

United Kingdom

Olea europaea

Common olive

Collar and trunk canker

Greece

Oryza sativa

Rice

Seedling disease

Japan

Persea americana

Avocado

Collar and trunk canker

Greece

Picea abies

Norway spruce

Root rot

New Zealand

Pinus spp.

Sugar pine, pine

Root rot; seedling damping-off

United States, New Zealand

Prunus spp.

Apricot; sweet, sour and Mahaleb cherry; common plum; almond; peach

Apoplexy; sudden wilt; root and crown rot; trunk canker

Australia, Greece, United States, Italy, New Zealand

Pseudotsuga menziesii

Douglas fir

Seedling rot; chlorosis; stunting

United States

Pyrus malus

Apple

Fruit rot

Japan

Rorippa spp.

Yellowcress

Root rot

United States

Rosa spp.

Rose

Stem blight

Japan

Rubus idaeus var. idaeus

Raspberry

Root rot

United Kingdom

Saccharum officinarum

Sugar cane

Seed-piece rot

United States

Senecio cruentus

Cineraria, groundsel

Root rot

United States

Solanum spp.

Eggplant, potato

Fruit rot; pink tuber rot

Japan, northern Ireland, United Kingdom, Canada, Australia

Spinacia oleracea

Spinach

Root rot

United States

Theobroma cacao

Cacao

Pod rot

Venezuela

Trifolium repens

White clover

Root rot

New Zealand, Australia

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

Barr, D. J. S. 1980. Heterothallic-like reaction in the large-oospore form of Phytophthora megasperma. Can. J. Plant Pathol. 2:116-118.

 

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 Virginia nursery irrigation water. Plant Health Progress doi:10.1094/PHP-2006-0621-01-RS.

 

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., Duncan, J. M., Wagels, G., and Brasier, C. M. 2000. A molecular phylogeny of Phytophthora and related oomycetes. Fungal Genet. Biol. 30:17-32.

 

Drechsler, C. 1931. A crown rot of hollyhocks caused by Phytophthora megasperma n. sp. J. Wash. Acad. Sci. 21:513-526.

 

Erwin, D. C., and Ribeiro, O. K. 1996. Phytophthora Diseases Worldwide. American Phytopathological Society, St. Paul, MN.

 

Faris, M. A., Sabo, F. E., Barr, D. J. S., and Lin, C. S. 1989. The systematics of Phytophthora sojae and P. megasperma. Can. J. Bot. 67:1442-1447.

 

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.

 

Gallegly, M. and Hong, C. 2008. Phytophthora: Identifying Species by Morphology and DNA Fingerprints. American Phytopathological Society Press, St. Paul, MN.

 

Ito, S., and Nagai, M., 1931. On the root disease of the seeds and seedlings of rice plants caused by some aquatic fungi. J. Fac. Agric. Hokkaido Univ. 32:45-69. (In Japanese)

 

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.

 

Ho, H. H. 1986. Notes on the heterothallic behavior of Phytophthora megasperma from alfalfa. Mycologia 78:306-309.

 

Kuan, T.-L., and Erwin, D. C. 1980. Formae speciales differentiation of Phytophthora megasperma isolates from soybean and alfalfa. Phytopathology 70:333-338.

 

Robertson, G. I., and Dance, H. M. 1971. The association of Phytophthora megasperma with crown rot of apple trees. N. Z. J. Agric. Res. 14:509-551.

 

Stamps, D. J., Newhook, F. J., Waterhouse, G. M., and Hall, G. S. 1990. Revised tabular key to the species of Phytophthora de Bary. Mycol. Pap. 162. CAB International, Wallingford, United Kingdom; Commonwealth Mycological Institute, Kew, Surrey, England.

 

Tompkins, C. M., Tucker, C. M., and Gardner, M. W. 1936. Phytophthora root rot of cauliflower. J. Agric. Res. 53:685-692.

 

Waterhouse, G. M. 1963. Key to the species of Phytophthora de Bary. Mycol. Pap. 92. CAB International, Wallingford, United Kingdom; Commonwealth Mycological Institute, Kew, Surrey, England.

 

Waterhouse, G. M., and Waterston, J. M. 1966. Phytophthora megasperma. CMI Descr. Pathog. Fungi Bact. 115.