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Powdery scab is a disease of
potato The potato () is a starchy tuberous vegetable native to the Americas that is consumed as a staple food in many parts of the world. Potatoes are underground stem tubers of the plant ''Solanum tuberosum'', a perennial in the nightshade famil ...
tuber Tubers are a type of enlarged structure that plants use as storage organs for nutrients, derived from stems or roots. Tubers help plants perennate (survive winter or dry months), provide energy and nutrients, and are a means of asexual reproduc ...
s. It is caused by the
cercozoa Cercozoa (now synonymised with Filosa) is a phylum of diverse single-celled eukaryotes. They lack shared morphological characteristics at the microscopic level, and are instead united by phylogeny, molecular phylogenies of rRNA and actin or Ubiqu ...
n ''Spongospora subterranea'' f. sp. ''subterranea'' and is widespread in potato growing countries.
Symptom Signs and symptoms are diagnostic indications of an illness, injury, or condition. Signs are objective and externally observable; symptoms are a person's reported subjective experiences. A sign for example may be a higher or lower temperature ...
s of powdery scab include small
lesion A lesion is any damage or abnormal change in the tissue of an organism, usually caused by injury or diseases. The term ''Lesion'' is derived from the Latin meaning "injury". Lesions may occur in both plants and animals. Types There is no de ...
s in the early stages of the
disease A disease is a particular abnormal condition that adversely affects the structure or function (biology), function of all or part of an organism and is not immediately due to any external injury. Diseases are often known to be medical condi ...
, progressing to raised
pustule A skin condition, also known as cutaneous condition, is any medical condition that affects the integumentary system—the organ system that encloses the body and includes skin, nails, and related muscle and glands. The major function of this ...
s containing a powdery mass. These can eventually rupture within the tuber periderm.Maldonado, M. Loreto Hernandez, Richard E. Falloon, Ruth C. Butler, Anthony J. Conner, and Simon R. Bulman. "''Resistance to Spongospora Subterranea Induced in Potato by the Elicitor β-aminobutyric Acid.''" Australasian Plant Pathol. Australasian Plant Pathology 44.4 (2015): 445-53. Web. 21 Oct. 2015. The powdery pustules contain resting spores that release anisokont
zoospore A zoospore is a motile asexual spore that uses a flagellum for locomotion in aqueous or moist environments. Also called a swarm spore, these spores are created by some protists, bacteria, and fungi to propagate themselves. Certain zoospores are ...
s (asexual spore with two unequal length
flagella A flagellum (; : flagella) (Latin for 'whip' or 'scourge') is a hair-like appendage that protrudes from certain plant and animal sperm cells, from fungal spores ( zoospores), and from a wide range of microorganisms to provide motility. Many pr ...
) to infect the root hairs of potatoes or tomatoes. Powdery scab is a cosmetic defect on tubers, which can result in the rejection of these potatoes. Potatoes which have been infected can be peeled to remove the infected skin and the remaining inside of the potato can be cooked and eaten.Dennis A. Johnson and Thomas F. Cummings. ''Effect of Powdery Scab Root Galls on Yield of Potato.'' (2015).Plant Disease 99:10, 1396-1403


Disease cycle

In general, not a lot is known about the life cycle of ''Spongospora subterranea'' f.sp ''subterranea'' (Sss). Most of the currently-proposed life cycle is based on that of ''
Plasmodiophora brassicae ''Plasmodiophora brassicae'' is a soilborne organism that is best known for its high economic impact on ''Brassica'' oil and vegetable crops. It infects 30 different Brassicaceae, cruciferous plants, which consists of up to 16 crop species, 9 or ...
'', a closely related and better-studied
protozoa Protozoa (: protozoan or protozoon; alternative plural: protozoans) are a polyphyletic group of single-celled eukaryotes, either free-living or parasitic, that feed on organic matter such as other microorganisms or organic debris. Historically ...
n. It has been proposed, due to this similarity, that there are two distinct stages that Sss can exist as; the asexual and sexual stages.


Asexual stage

A
zoospore A zoospore is a motile asexual spore that uses a flagellum for locomotion in aqueous or moist environments. Also called a swarm spore, these spores are created by some protists, bacteria, and fungi to propagate themselves. Certain zoospores are ...
infects root tissue and becomes a uninucleate
plasmodium ''Plasmodium'' is a genus of unicellular eukaryotes that are obligate parasites of vertebrates and insects. The life cycles of ''Plasmodium'' species involve development in a Hematophagy, blood-feeding insect host (biology), host which then inj ...
. This plasmodium undergoes
mitotic Mitosis () is a part of the cell cycle in eukaryotic cells in which replicated chromosomes are separated into two new nuclei. Cell division by mitosis is an equational division which gives rise to genetically identical cells in which the t ...
nuclear division (creates many nuclei within a single cell) and turns into a
multinucleate Multinucleate cells (also known as multinucleated cells or polynuclear cells) are eukaryotic cells that have more than one nucleus, i.e., multiple nuclei share one common cytoplasm. Mitosis in multinucleate cells can occur either in a coordinate ...
plasmodium. Then, the multinucleate plasmodium forms zoosporangium, which eventually release more zoospores. This process can happen relatively quickly and can act as an important source of secondary inoculum within a field.


Sexual stage

This stage follows a similar pattern to the asexual stage, but with a few exceptions. It is hypothesized that two zoospores fuse together to form a dikaryotic zoospore (with two separate haploid nuclei, n+n) and then infect the roots. Once the infection occurs, the dikaryotic zoospore develops into a binucleate plasmodium (one pair on nuclei, n+n). Similar to the asexual stage, this plasmodium will also replicate its nucleus to create a multinucleate plasmodium (many pairs of nuclei, n+n). The second main different between stages occurs here. The pairs of nuclei (n+n) will fuse by
karyogamy Karyogamy is the final step in the process of fusing together two haploid eukaryotic cells, and refers specifically to the fusion of the two cell nucleus, nuclei. Before karyogamy, each haploid cell has one complete copy of the organism's genome. ...
, and the plasmodium will quickly divide into numerous resting spores within a sporosori (spore sack, alternatively called cystosori). These resting spores have three-layered walls and are extremely resistant to the environment, allowing them to persist in the soil for longer than 10 years. As a reminder, most of the life cycle is still unclear. However, the presence of zoospores, plasmodia, zoosporangia, and resting spores have been observed in the field and lab. The
ploidy Ploidy () is the number of complete sets of chromosomes in a cell, and hence the number of possible alleles for autosomal and pseudoautosomal genes. Here ''sets of chromosomes'' refers to the number of maternal and paternal chromosome copies, ...
levels and karyogamy events are only theorized and have yet to be proven.


Environment

''Spongospora subterranea'' pathogenesis is most effective in cool, damp environments, such as northern Britain, the Columbia Basin of south-central Washington, and north-central Oregon. The environmental condition is particularly critical during the release of infective agents (zoospores) into the soil-environment . Upon release from resting spores, zoospores require moisture to swim towards the host tuber or roots. One study, found powdery scab was significantly more common on plants grown in constant dampness compared to plants grown with varying moisture levels. In this same study it was concluded that disease risk was related more to the environment, or moisture level, than the level of inoculum present. Inoculum may be present but not able to disperse due to environmental conditions, and therefore does not reach host tissue to infect. Other environmental factors that affect ''Spongospora subterranea'' infection are directly related to agronomic practices. Increased use of fertilizers containing
nitrate Nitrate is a polyatomic ion with the chemical formula . salt (chemistry), Salts containing this ion are called nitrates. Nitrates are common components of fertilizers and explosives. Almost all inorganic nitrates are solubility, soluble in wa ...
or ammonium nitrogen increase the incidence and severity of powdery scab. It is thought that the fertilization increases root growth, and thus provides more tissue for infection and disease cycling to occur. Also, reduced
cellulose Cellulose is an organic compound with the chemical formula, formula , a polysaccharide consisting of a linear chain of several hundred to many thousands of glycosidic bond, β(1→4) linked glucose, D-glucose units. Cellulose is an important s ...
within the cell walls caused by excess nitrogen may increase susceptibility of host to infection. It is apparent that the environment can directly affect both the host susceptibility and the dispersal of the pathogen ultimately setting the pace for the disease cycle.


Pathogenesis

''S. subterranea'' is an obligate parasite
phytomyxea The Phytomyxea are a class (biology), class of parasites that are cosmopolitan, obligate biotrophic protist parasites of plants, diatoms, oomycetes and brown algae. They are divided into the orders Plasmodiophorida (International Code of Zoolog ...
that infects the below ground structures of the host. Infection leads to
hypertrophy Hypertrophy is the increase in the volume of an organ or tissue due to the enlargement of its component cells. It is distinguished from hyperplasia, in which the cells remain approximately the same size but increase in number. Although hypertro ...
and
hyperplasia Hyperplasia (from ancient Greek ὑπέρ ''huper'' 'over' + πλάσις ''plasis'' 'formation'), or hypergenesis, is an enlargement of an organ or tissue caused by an increase in the amount of Tissue (biology), organic tissue that results from ...
of the host cells and eventual bursting. However, the mechanism behind this is still unknown. Zoospores infect the
root hair Soil biology Root hairs or absorbent hairs, are outgrowths of epidermal cells, specialized cells at the tip of a plant root. They are lateral extensions of a single cell and are only rarely branched. They are found in the region of maturation, of ...
s by attaching to the outer surface, encysting, and then penetrating the
epidermis The epidermis is the outermost of the three layers that comprise the skin, the inner layers being the dermis and Subcutaneous tissue, hypodermis. The epidermal layer provides a barrier to infection from environmental pathogens and regulates the ...
through
lenticel A lenticel is a porous tissue consisting of cells with large intercellular spaces in the periderm of the secondarily thickened organs and the Bark (botany), bark of woody stems and roots of gymnosperms and dicotyledonous flowering plants. It func ...
s and
stoma In botany, a stoma (: stomata, from Greek language, Greek ''στόμα'', "mouth"), also called a stomate (: stomates), is a pore found in the Epidermis (botany), epidermis of leaves, stems, and other organs, that controls the rate of gas exc ...
ta. Once inside, the multinucleate plasmodium divides to spread and produce more zoospores. The plasmodium causes the infected host cells to multiply rapidly and enlarge into a gall. This rapid multiplication also produces uninucleate cells that aggregate together as sporosori. The sporosori look like a powdery mass within the gall, which gives this disease its name. Eventually the gall swells and bursts out the epidermis of the tuber, releasing the spores back into the soil. Gall severity depends on inoculum level, environment, and potato skin type. Infection is most prevalent in the early stages of tuber formation while the potato tissue is unsuberized. But, infection can occur at all stages on development. White and red skinned potatoes and highly susceptible while russet skinned are somewhat resistant. Russet skin is thicker and has higher levels of the LOX protein which is used as a marker for resistance. There is little known about variation and sexual recombination within ''S. subterranea,'' therefore high priority is given to researching the variations within potato cultivars for researching host/pathogen relationships and management.


Importance

Powdery Scab has important implications for commercial farming. Not only does the pathogen itself cause harm, but the pathogen is also a vector for potato mop-top virus, another plant pathogen. As a result, its presence greatly threatens potato marketability for farmers. The burst pustules can also act as a wound for other fungi to infect, such as '' Phytophthora erythroseptica'' and ''
Phytophthora infestans ''Phytophthora infestans'' is an oomycete or Oomycete, water mold, a fungus-like microorganism that causes the serious potato and tomato disease known as late blight or potato blight. Early blight, caused by ''Alternaria solani'', is also often c ...
''. Thus, tubers with powdery scab can have increased incidences of other devastating diseases, including pink rot, dry rot, black dot, and late blight. Potato tubers will form powdery scab pustules that inhibit their ability to be sold. Many markets decline to buy potatoes with ugly scarring even if they are safe to eat. Research has not yet found an effective way to peel the scabs without damaging the potato. Potatoes that are rejected for sale create a large financial burden on farmers. Additionally, because soil borne inoculum can survive for years as spores, the pathogen is very difficult to eliminate once present. In Great Britain a recent Potato Council funded diagnostic project discovered that as much as 82% of fields tested positive for soil inoculum.


Management

''S. subterranea'' currently has no effective chemical controls. Therefore, other cultural management techniques must be used. Using certified clean seeds and planting in fields that have been historically healthy is the best form of control. These methods may prevent infestation from resting spores. Since infection is promoted by cool soil temperatures and high soil moisture, delayed planting can also help reduce negative effects of the pathogen. Delayed planting reduces the growth period in cooler soils subsequently decreasing germination of the spores. One limitation to this method is an additional decrease of early market yield. Pre-planting chemigation with metam sodium can reduce the propagules of the pathogen."How to Manage Pests." ''UC IPM: UC Management Guidelines for Powdery Scab on Potato''. N.p., n.d. Web. 19 Oct. 2015. Other common means of control include using resistant potatoes and crop rotations. Several cultivars of resistant potatoes include Granola, Nicola, Ditta, and Gladiator. Because soil-borne inoculum can survive for many years,
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. This practice reduces the reliance of crops on one set of nutrients, pest and weed pressure, along with the pro ...
s should involve alternate species that will promote a partial life cycle of the pathogen. This way the
zoospore A zoospore is a motile asexual spore that uses a flagellum for locomotion in aqueous or moist environments. Also called a swarm spore, these spores are created by some protists, bacteria, and fungi to propagate themselves. Certain zoospores are ...
s will germinate without producing new spores. Researchers have investigated the use of beta-aminobutyric acid (BABA) in promoting potato resistance. BABA triggers a plants
systemic acquired resistance Systemic acquired resistance (SAR) is a "whole-plant" resistance response that occurs following an earlier localized exposure to a pathogen. SAR is analogous to the innate immune system found in animals, and although there are many shared aspects b ...
(SAR), a natural plant defense mechanism. When potatoes are inoculated with BABA and then later inoculated with the pathogen, ''S. subterranea,'' they exhibit overall reduction in disease. While pathogen reduction has been experimentally supported, further experimentation needs to be performed. Genomic data inclusive a genome draft became recently available, that might hold information that can be used to improve disease management.


References


External links


Powdery scab photos
from Cornell University
Resistance to Powdery Scab in Potato
from the US Department of Agriculture {{Authority control Potato diseases Endomyxa Parasitic rhizaria Forma specialis taxa