Phytophthora Tabaci
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''Phytophthora nicotianae'' or black shank is an
oomycete Oomycota forms a distinct phylogenetic lineage of fungus-like eukaryotic microorganisms, called oomycetes (). They are filamentous and heterotrophic, and can reproduce both sexually and asexually. Sexual reproduction of an oospore is the resul ...
belonging to the order Peronosprales and family
Peronosporaceae Peronosporaceae are a family of water moulds that contains 21 genera, comprising more than 600 species. Most of them are called downy mildews. Peronosporaceae are obligate biotrophic plant pathogens. They parasitise their host plants as an ...
.


Hosts and symptoms

''Phytophthora nicotianae'' has a broad host range comprising 255 genera from 90
families Family (from la, familia) is a group of people related either by consanguinity (by recognized birth) or affinity (by marriage or other relationship). The purpose of the family is to maintain the well-being of its members and of society. Ideal ...
. Hosts include
tobacco Tobacco is the common name of several plants in the genus '' Nicotiana'' of the family Solanaceae, and the general term for any product prepared from the cured leaves of these plants. More than 70 species of tobacco are known, but the ...
,
onion An onion (''Allium cepa'' L., from Latin ''cepa'' meaning "onion"), also known as the bulb onion or common onion, is a vegetable that is the most widely cultivated species of the genus ''Allium''. The shallot is a botanical variety of the onio ...
,
tomato The tomato is the edible berry of the plant ''Solanum lycopersicum'', commonly known as the tomato plant. The species originated in western South America, Mexico, and Central America. The Mexican Nahuatl word gave rise to the Spanish word ...
,
ornamentals Ornamental plants or garden plants are plants that are primarily grown for their beauty but also for qualities such as scent or how they shape physical space. Many flowering plants and garden varieties tend to be specially bred cultivars that i ...
,
cotton Cotton is a soft, fluffy staple fiber that grows in a boll, or protective case, around the seeds of the cotton plants of the genus '' Gossypium'' in the mallow family Malvaceae. The fiber is almost pure cellulose, and can contain minor pe ...
,
pepper Pepper or peppers may refer to: Food and spice * Piperaceae or the pepper family, a large family of flowering plant ** Black pepper * ''Capsicum'' or pepper, a genus of flowering plants in the nightshade family Solanaceae ** Bell pepper ** Chili ...
, and
citrus ''Citrus'' is a genus of flowering trees and shrubs in the rue family, Rutaceae. Plants in the genus produce citrus fruits, including important crops such as oranges, lemons, grapefruits, pomelos, and limes. The genus ''Citrus'' is native to ...
plants. This pathogen can cause root rot, crown rot, fruit rot, leaf infection, and stem infection. Root rot symptoms are observed on tobacco, poinsettia, tomato,
pineapple The pineapple (''Ananas comosus'') is a tropical plant with an edible fruit; it is the most economically significant plant in the family Bromeliaceae. The pineapple is indigenous to South America, where it has been cultivated for many centuri ...
,
watermelon Watermelon (''Citrullus lanatus'') is a flowering plant species of the Cucurbitaceae family and the name of its edible fruit. A scrambling and trailing vine-like plant, it is a highly cultivated fruit worldwide, with more than 1,000 varie ...
, and as well as
African violet ''Streptocarpus'' sect. ''Saintpaulia'' is a section within ''Streptocarpus'' subgenus ''Streptocarpella'' consisting of about ten species of herbaceous perennial flowering plants in the family Gesneriaceae, native to Tanzania and adjacent sout ...
. Fruit rots occur on tomato, papaya, and eggplant. Onion shows a leaf and stem infection. In tobacco Black Shank affects the roots and basal stem area, but all parts of the plant can become infected.
Damping off Damping off (or damping-off) is a horticultural disease or condition, caused by several different pathogens that kill or weaken seeds or seedlings before or after they germinate. It is most prevalent in wet and cool conditions. Symptoms There ar ...
symptoms can be observed in young seedlings. The first above ground symptom that will be observed is the wilting of plants, which leads to stunting. Roots will be blackened and decayed. In final stages of the disease the stem begins to turn black, hence the name Black Shank. As this happens, tobacco leaves turn brown and become not marketable. Another symptom is disk-like appearance of the
pith Pith, or medulla, is a tissue in the stems of vascular plants. Pith is composed of soft, spongy parenchyma cells, which in some cases can store starch. In eudicotyledons, pith is located in the center of the stem. In monocotyledons, it ext ...
, although this is not a definitive symptom as it may also be the result of
lightning Lightning is a naturally occurring electrostatic discharge during which two electrically charged regions, both in the atmosphere or with one on the ground, temporarily neutralize themselves, causing the instantaneous release of an avera ...
strikes. On
onion An onion (''Allium cepa'' L., from Latin ''cepa'' meaning "onion"), also known as the bulb onion or common onion, is a vegetable that is the most widely cultivated species of the genus ''Allium''. The shallot is a botanical variety of the onio ...
it causes the disease known as ''Phytophthora'' neck and bulb rot. Different stages of onion may be affected. Initially, tips of newly infected plants start to yellow and dry followed by softening of the "neck" of the plants that eventually fall over. Infected leaves may show grey lesions. Roots may become necrotic in late disease.


Disease cycle

Black Shank is a polycyclic soil borne disease, with the possibility of multiple disease cycles per growing season occurring from May to October. There are important structures this pathogen uses in its disease cycle.
Chlamydospore A chlamydospore is the thick-walled large resting spore of several kinds of fungi, including Ascomycota such as '' Candida'', Basidiomycota such as '' Panus'', and various Mortierellales species. It is the life-stage which survives in unfavourable ...
s are produced asexually and serve as long lived resting structures, surviving from four to six years. Chlamydospores are the primary survival structure, the primary inoculum, and are usually produced in abundance. These spores germinate in warm and moist soil to produce a
germ tube A germ tube is an outgrowth produced by spores of spore-releasing fungi during germination. The germ tube differentiates, grows, and develops by mitosis to create somatic hyphae.C.J. Alexopolous, Charles W. Mims, M. Blackwell, ''Introductory My ...
that infects plants or produces a sporangium. Another asexual structure and secondary inoculum, appearing ovoid, pear, or spherical in shape are called sporangium. These spores are produced and can either germinate directly or release motile
zoospore A zoospore is a motile asexual spore that uses a flagellum for locomotion. Also called a swarm spore, these spores are created by some protists, bacteria, and fungi to propagate themselves. Diversity Flagella types Zoospores may possess one or ...
s within 24 hours of inoculation with the right conditions. Zoospores are kidney shaped with an anterior tinsel
flagellum A flagellum (; ) is a hairlike appendage that protrudes from certain plant and animal sperm cells, and from a wide range of microorganisms to provide motility. Many protists with flagella are termed as flagellates. A microorganism may have f ...
and a posterior whip like flagellum that helps to navigate toward root tips were infection occurs. Black Shank needs water for germination and movement because zoospores swim through soil pores and standing water. Splashing water from rain or irrigation can infect healthy plant leaves leading to more repeating secondary cycles. Zoospores move toward nutrient gradients around root tips and host wounds. Once the root surface is contacted, zoospores encyst and a germ tube will emerge penetrating the epidermis. Infection leads to systemic rotting of the root system and wilting and
chlorosis In botany, chlorosis is a condition in which leaves produce insufficient chlorophyll. As chlorophyll is responsible for the green color of leaves, chlorotic leaves are pale, yellow, or yellow-white. The affected plant has little or no ability to ...
in the leaves. Another structure called hyphae is colorless, transparent, and coenocytic, but colonies may yellow with age. Also, there is much morphological variation in colony type with different isolates of ''P. nicotianae'' and the growth may differ when grown on different media. The hyphae are
heterothallic Heterothallic species have sexes that reside in different individuals. The term is applied particularly to distinguish heterothallic fungi, which require two compatible partners to produce sexual spores, from homothallic ones, which are capable ...
and require two
mating type Mating types are the microorganism equivalent to sexes in multicellular lifeforms and are thought to be the ancestor to distinct sexes. They also occur in macro-organisms such as fungi. Definition Mating types are the microorganism equivalent to ...
s to produce
oospore An oospore is a thick-walled sexual spore that develops from a fertilized oosphere in some algae, fungi, and oomycetes. They are believed to have evolved either through the fusion of two species or the chemically-induced stimulation of mycelia ...
s, the sexual survival structure. Many fields only contain one mating type, so the zoospores rarely germinate and rarely cause epidemics.


Environment

This pathogen thrives in temperatures ranging from . Disease is prominent in many agricultural productive regions and therefore is a major host to many warm environment crops. Black Shank needs water for germination and movement. Saturated soil optimizes disease spread because water is used for dissemination of motile zoospores and sporangia. Low-lying areas of the soil that remain wet for prolonged periods of time will have more disease. Splashing water from rain or
irrigation Irrigation (also referred to as watering) is the practice of applying controlled amounts of water to land to help grow crops, landscape plants, and lawns. Irrigation has been a key aspect of agriculture for over 5,000 years and has been devel ...
can infect healthy plant leaves leading to more repeating secondary cycles. Soils that are not saturated will lead to little to no disease development, so water management is important. Optimum soil pH for development is between 6 and 7. Levels of
calcium Calcium is a chemical element with the symbol Ca and atomic number 20. As an alkaline earth metal, calcium is a reactive metal that forms a dark oxide-nitride layer when exposed to air. Its physical and chemical properties are most similar t ...
and
magnesium Magnesium is a chemical element with the symbol Mg and atomic number 12. It is a shiny gray metal having a low density, low melting point and high chemical reactivity. Like the other alkaline earth metals (group 2 of the periodic ta ...
in the soils can affect disease progress.


Management

Several kinds of management exist for the prevention and suppression of disease. A cultural method that can be effective in preventing disease is sanitation. Equipment should be cleaned after use in infested fields so the disease does not spread into uninfested fields. To disrupt chlamydospore germination crops should be grown in drained disease free soil. Also, avoid transplanting without thorough knowledge of the transplant. To limit spread of structures limit traffic in infected fields and always clean after exposure. Disease is favored by pH values greater than 6.2, so lowering the pH is an effective method for preventing germination. pH management can be difficult because tobacco cannot survive in very low pH soils. Soil pH 5.5 to 6 allow successful growth of tobacco and control of disease. The cultural control,
Crop rotation Crop rotation is the practice of growing a series of different types of crops in the same area across a sequence of growing seasons. It reduces reliance on one set of nutrients, pest and weed pressure, and the probability of developing resistant ...
, is very effective at limiting disease. The longer an infected field is planted in a crop other than the initial infected crop, the lower the population will become. A minimum three-year rotation is recommended. Crop rotation is recommended in combination with resistant varieties as genetic controls.
Burley Tobacco Burley tobacco is a light air-cured tobacco used primarily for cigarette production. In the United States it is produced in an eight-state belt with approximately 70% produced in Kentucky. Tennessee produces approximately 20%, with smaller amoun ...
, Burley Tobacco hybrids, and Dark Tobacco are varieties of tobacco that are resistant to Black Shank. Resistance however is not reliable because a single variety has resistance to only a few races of Black Shank. Finding new lines of resistance is becoming increasingly important due to new discovered resistant races of the pathogen. Chemical control is most successful if used with resistant varieties.
Metalaxyl Metalaxyl is an acylalanine fungicide with systemic function. Its chemical name is methyl N-(methoxyacetyl)-N-(2,6-xylyl)-DL-alaninate. It can be used to control ''Pythium'' in a number of vegetable crops, and ''Phytophthora'' in peas. Metalaxyl- ...
or mefenoxam are chemistries used to control ''Phytophthora nicotianae''. Ridomil Gold is an example a systemic pesticide with a Metalaxyl chemistry. Mefenoxam is twice as active as metalaxyl, but they both have the same mode of action. Successful chemical control is difficult because we are limited to these two chemistries that are basically identical. A study by A. S. Csinos and P. F. Bertrand found out at a rate of 3.36 kg/ha would not inhibit many of the common races used in their study. Overall, from their study they observed that Black Shank severity was increasing in
Georgia Georgia most commonly refers to: * Georgia (country), a country in the Caucasus region of Eurasia * Georgia (U.S. state), a state in the Southeast United States Georgia may also refer to: Places Historical states and entities * Related to the ...
due to Metalaxyl sensitivity and resistant races of Black Shank.


Importance

''Phytophthora nicotianae'' has a wide host range, affecting agriculture rich areas all over the world. In the
United States The United States of America (U.S.A. or USA), commonly known as the United States (U.S. or US) or America, is a country primarily located in North America. It consists of 50 states, a federal district, five major unincorporated territori ...
this is a major pathogen of ornamentals, tobacco, and tomato. Black Shank is one of the most damaging and far reaching diseases of tobacco. In 1896, Black Shank was first described in
Indonesia Indonesia, officially the Republic of Indonesia, is a country in Southeast Asia and Oceania between the Indian and Pacific oceans. It consists of over 17,000 islands, including Sumatra, Java, Sulawesi, and parts of Borneo and New Guine ...
by Van Breda de Haan. Disease was observed near Georgia in 1915 and reached major tobacco growing areas of
Kentucky Kentucky ( , ), officially the Commonwealth of Kentucky, is a state in the Southeastern region of the United States and one of the states of the Upper South. It borders Illinois, Indiana, and Ohio to the north; West Virginia and Virginia ...
and
North Carolina North Carolina () is a state in the Southeastern region of the United States. The state is the 28th largest and 9th-most populous of the United States. It is bordered by Virginia to the north, the Atlantic Ocean to the east, Georgia and ...
in the 1930s and 1940s. In North Carolina black shank can be found in every county that grows flue-cured tobacco and currently causes statewide losses of 1 to 2.5 percent per year. This pathogen thrives in warm climates, so it is destructive on crops grown in these areas. During favorable conditions, new generations of spores can be produced every 72 hours, so if this disease is not managed well it can be very destructive. Susceptible
cultivar A cultivar is a type of cultivated plant that people have selected for desired traits and when propagated retain those traits. Methods used to propagate cultivars include: division, root and stem cuttings, offsets, grafting, tissue culture ...
s in the right conditions can reach losses of 100 percent, because infected plants do not recover. Less than one
propagule In biology, a propagule is any material that functions in propagating an organism to the next stage in its life cycle, such as by dispersal. The propagule is usually distinct in form from the parent organism. Propagules are produced by organisms ...
per gram of soil can lead to an
epidemic An epidemic (from Greek ἐπί ''epi'' "upon or above" and δῆμος ''demos'' "people") is the rapid spread of disease to a large number of patients among a given population within an area in a short period of time. Epidemics of infectious ...
.


Pathogenesis

This pathogen causes secondary cycles of disease by mode of zoospores. Zoospores interact with the host by sensing and moving toward the nutrient gradients near the root tip and wounds of the plant. Without this means of sensing entry points there would be no secondary cycles of disease. Zoospores, chlamydospores, and sporangia produce a germ tube that directly penetrates the epidermis of the plant. Without this penetration device the pathogen would not be able to infect the plant. The pathogen interferes with transport by infecting the roots. Typically hyphae can be seen in the pith and cause blackening and necrosis. Infection can proceed rapidly once the pathogen has made an entrance into the plant. Once established, further reproduction of both chlamydospores and sporangia will occur within host tissues, amplifying the spread of disease within the host plant and spreading out into nearby plants. Upon death of the host, the decomposing infected tissues will release the pathogen back into the soil, in the form of chlamydospores and zoospores. A resting spore, the chlamydospores are capable of surviving in the soil for years, but it has been noted that cold winters cause an inhibitory effect on the survival rate. This results in less black shank infections where tobacco is grown in cooler, more northern climates. The action of ''P. nicotianae'' is amplified by the presence of
root-knot nematode Root-knot nematodes are plant-parasitic nematodes from the genus ''Meloidogyne''. They exist in soil in areas with hot climates or short winters. About 2000 plants worldwide are susceptible to infection by root-knot nematodes and they cause appr ...
s, which through their own feeding habits, assist the pathogen in finding an entrance to the host. This pathogen synergy with root-knot nematodes has the ability to overcome much of the resistance of cultivars especially bred for ''P. nicotianae'' resistance.


References

{{DEFAULTSORT:Phytophthora Nicotianae nicotianae Species described in 1896 Water mould plant pathogens and diseases Vegetable diseases