Powdery scab
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Powdery scab is a disease of
potato The potato is a starchy food, a tuber of the plant ''Solanum tuberosum'' and is a root vegetable native to the Americas. The plant is a perennial in the nightshade family Solanaceae. Wild potato species can be found from the southern Unit ...
tuber Tubers are a type of enlarged structure used as storage organs for nutrients in some plants. They are used for the plant's perennation (survival of the winter or dry months), to provide energy and nutrients for regrowth during the next growin ...
s. It is caused by the cercozoan ''Spongospora subterranea'' f. sp. ''subterranea'' and is widespread in potato growing countries. Symptoms of powdery scab include small
lesion A lesion is any damage or abnormal change in the tissue of an organism, usually caused by disease or trauma. ''Lesion'' is derived from the Latin "injury". Lesions may occur in plants as well as animals. Types There is no designated classif ...
s in the early stages of the
disease A disease is a particular abnormal condition that negatively affects the structure or function of all or part of an organism, and that is not immediately due to any external injury. Diseases are often known to be medical conditions that a ...
, 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 ...
s containing a powdery mass. These can eventually rupture within the tuber
periderm Bark is the outermost layers of stems and roots of woody plants. Plants with bark include trees, woody vines, and shrubs. Bark refers to all the tissues outside the vascular cambium and is a nontechnical term. It overlays the wood and consist ...
.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. 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 (asexual spore with two unequal length flagella) 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'' is a genus in the phylum of cercozoa in the class Phytomyxea. It includes the species ''Plasmodiophora brassicae'', which causes the disease cabbage clubroot Clubroot is a common disease of cabbages, broccoli, caulifl ...
'', a closely related and better-studied protozoan. 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. 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 ...
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 blood-feeding insect host which then injects parasites into a ver ...
. This plasmodium undergoes
mitotic In cell biology, mitosis () is a part of the cell cycle in which replicated chromosomes are separated into two new nuclei. Cell division by mitosis gives rise to genetically identical cells in which the total number of chromosomes is maintai ...
nuclear division (creates many nuclei within a single cell) and turns into a
multinucleate Multinucleate cells (also known as multinucleated or polynuclear cells) are eukaryotic cells that have more than one nucleus per cell, i.e., multiple nuclei share one common cytoplasm. Mitosis in multinucleate cells can occur either in a coordina ...
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, and the plasmodium will quickly divide into numerous
resting spore A resting spore is a resistant cell, used to survive adverse environmental conditions. Resting spore is a term commonly applied to both diatoms and fungi. In fungi A resting spore can be a spore created by fungi which is thickly encysted (has a ...
s 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 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 Agricultural economics is an applied field of economics concerned with the application of economic theory in optimizing the production and distribution of food and fiber products. Agricultural economics began as a branch of economics that spe ...
practices. Increased use of fertilizers containing nitrate 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 formula , a polysaccharide consisting of a linear chain of several hundred to many thousands of β(1→4) linked D-glucose units. Cellulose is an important structural component of the primary cell w ...
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 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.Updated by Linda J. ...
and hyperplasia of the host cells and eventual bursting. However, the mechanism behind this is still unknown. Zoospores infect the
root hair Root hair, 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 the root. Root ...
s by attaching to the outer surface, encysting, and then penetrating the epidermis through lenticels and stomata. 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 ''Potato mop-top virus (PMTV)'' is a plant pathogenic virus transmitted through the vector ''Spongospora subterranea'' that affects potatoes. PMTV belongs to family of ''Virgaviridae'', and the genus ''Pomovirus'' (Potato mop-top virus). The viru ...
, 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 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 called "p ...
''. Thus, tubers with powdery scab can have increased incidences of other devastating diseases, including pink rot, dry rot, black dot, and
late blight ''Phytophthora infestans'' is an oomycete or 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 called " ...
. 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 Metam sodium is an organosulfur compound with the formula . The compound is a sodium salt of a dithiocarbamate. The compound exists as a colorless dihydrate, but most commonly it is encountered as an aqueous solution. It is used as a soil fumig ...
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. It reduces reliance on one set of nutrients, pest and weed pressure, and the probability of developing resistant ...
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. 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 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 ...
(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 {{Taxonbar, from=Q1431083 Potato diseases Endomyxa Parasitic rhizaria Forma specialis taxa