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''Pythium sulcatum'' is a
chromalveolate Chromalveolata was a eukaryote supergroup present in a major classification of 2005, then regarded as one of the six major groups within the eukaryotes. It was a refinement of the kingdom Chromista, first proposed by Thomas Cavalier-Smith in ...
plant pathogen Plant pathology (also phytopathology) is the scientific study of diseases in plants caused by pathogens (infectious organisms) and environmental conditions (physiological factors). Organisms that cause infectious disease include fungi, oomy ...
infecting
carrot The carrot ('' Daucus carota'' subsp. ''sativus'') is a root vegetable, typically orange in color, though purple, black, red, white, and yellow cultivars exist, all of which are domesticated forms of the wild carrot, ''Daucus carota'', na ...
s. Because this organism was once thought to be a type of
fungus A fungus ( : fungi or funguses) is any member of the group of eukaryotic organisms that includes microorganisms such as yeasts and molds, as well as the more familiar mushrooms. These organisms are classified as a kingdom, separately fr ...
, it is still often treated as such.


Host and symptoms

''Pythium sulcatum'' causes diseases predominately on members of the apiaceous family, the most common being the carrot. This pathogen causes cavity spots which are sunken, spherical to elliptical oriented, brown-black spots across the breadth of the surface of the carrot. These cavity spots are normally 1-10 mm in width and are often surrounded by a pale halo. The lesions enlarge as the roots mature, often most prominent around the time the crop is harvested. This pathogen is believed to be native of the apiaceous family that eventually became pathogenic to carrots. ''P. sulcatum'' is also seen to cause
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 ...
, taproot dieback,
root rot Root rot is a condition in which anoxic conditions in the soil or potting media around the roots of a plant cause them to rot. This occurs due to excessive standing water around the roots. It is found in both indoor and outdoor plants, although ...
ting, stunting, and forking of carrots, with cavity spots being the most common of these diseases. Because carrots are a
root vegetable Root vegetables are underground plant parts eaten by humans as food. Although botany distinguishes true roots (such as taproots and tuberous roots) from non-roots (such as bulbs, corms, rhizomes, and tubers, although some contain both hypocoty ...
the cavity spots are only apparent after the carrot is harvested and washed.


Environment

''Pythium sulcatum'' can cause disease in both
mineral In geology and mineralogy, a mineral or mineral species is, broadly speaking, a solid chemical compound with a fairly well-defined chemical composition and a specific crystal structure that occurs naturally in pure form.John P. Rafferty, ed. (2 ...
and
organic soil Soil, also commonly referred to as earth or dirt, is a mixture of organic matter, minerals, gases, liquids, and organisms that together support life. Some scientific definitions distinguish ''dirt'' from ''soil'' by restricting the former term ...
s. The pathogen prefers wet to moist soils with a relatively acidic pH content, as a pH of 6.8 is ideal. However, the pathogen has been seen to grow in acidic pH of 3-5 and basic pH of 9-10. ''P. sulcatum'' prefers a relatively warm temperature, with 20°C–28°C being the most common, 25°C being optimal. This is why the pathogen is most severe in summer and autumn-harvested crops. The pathogen overwinters in the form of an
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 ...
, which can survive several years in the soil and inoculates the host when temperatures become optimal. In heavy moisture and wet soil, the pathogen can also release zoospores that can further increase its population up to 1000-fold. Flooding of soil by heavy rains as well as poorly drained soils have been shown to increase cavity spot disease development.


Pathogenesis

''Pythium sulcatum'' 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 ...
that is slow growing, aggressively virulent, and consistently pathogenic. Pathogenesis begins with the germination of resting spores (
oogonia An oogonium (plural oogonia) is a small diploid cell which, upon maturation, forms a primordial follicle in a female fetus or the female (haploid or diploid) gametangium of certain thallophytes. In the mammalian fetus Oogonia are formed in l ...
and
hypha A hypha (; ) is a long, branching, filamentous structure of a fungus, oomycete, or actinobacterium. In most fungi, hyphae are the main mode of vegetative growth, and are collectively called a mycelium. Structure A hypha consists of one or ...
l swellings), that occurs quickly in response to
root exudate Plant root exudates are fluids emitted through the roots of plants. These secretions influence the rhizosphere around the roots to inhibit harmful microbes and promote the grow of self and kin plants. Plant root systems can grow to be complex du ...
s. This causes direct infection through the unwounded surface of the root. The lesions form under the intact
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 ...
(corky outer layer of the root) and later ruptures causing dark, elongated lesions to develop. This typically occurs randomly on the root or may be more dense on the upper half. ''P. sulcatum'' is associated with its ability to produce a wide array of
cell wall A cell wall is a structural layer surrounding some types of cells, just outside the cell membrane. It can be tough, flexible, and sometimes rigid. It provides the cell with both structural support and protection, and also acts as a filtering mec ...
degrading enzymes with significantly high enzymatic activity. These enzymes are:
polygalacturonase Endo-polygalacturonase (EC 3.2.1.15, pectin depolymerase, pectolase, pectin hydrolase, and poly-α-1,4-galacturonide glycanohydrolase; systematic name (1→4)-α-D-galacturonan glycanohydrolase (endo-cleaving)) is an enzyme that hydrolyzes the α- ...
,
pectin lyase Pectin lyase (), also known as pectolyase, is a naturally occurring pectinase, a type of enzyme that degrades pectin. It is produced commercially for the food industry from fungi and used to destroy residual fruit starch, known as pectin, in wine ...
, lactate lyase,
cellulase Cellulase (EC 3.2.1.4; systematic name 4-β-D-glucan 4-glucanohydrolase) is any of several enzymes produced chiefly by fungi, bacteria, and protozoans that catalyze cellulolysis, the decomposition of cellulose and of some related polysaccha ...
, and pectin methylesterase. The digestion of the host cell wall and tissue maceration through the cell-wall degrading enzymes are crucial aspects of penetration and colonization of the carrot tissues in cavity spot pathogenesis. ''P. sulcatum'' first begins to secrete polygalacturonase and pectin methylesterase which begins to degrade the
pectin Pectin ( grc, πηκτικός ': "congealed" and "curdled") is a heteropolysaccharide, a structural acid contained in the primary lamella, in the middle lamella, and in the cell walls of terrestrial plants. The principal, chemical component o ...
in the plant cell’s cell wall. The pathogen then begins to secrete the three-remaining cell-wall degrading enzymes (pectin lyase,
pectate lyase Pectate lyase () is an enzyme involved in the maceration and soft rotting of plant tissue. Pectate lyase is responsible for the eliminative cleavage of pectate, yielding oligosaccharides with 4-deoxy-α-D-mann-4-enuronosyl groups at their non-re ...
, and cellulase) as well as ß-1,4-glucanase and
xylanase Endo-1,4-β-xylanase (EC 3.2.1.8, systematic name 4-β-D-xylan xylanohydrolase) is any of a class of enzymes that degrade the linear polysaccharide xylan into xylose, thus breaking down hemicellulose, one of the major components of plant cel ...
which further degrade the pectin in the plant cell’s cell wall into
oligomer In chemistry and biochemistry, an oligomer () is a molecule that consists of a few repeating units which could be derived, actually or conceptually, from smaller molecules, monomers.Quote: ''Oligomer molecule: A molecule of intermediate relati ...
s. This is the pathogenic pathway of how ''P. sulcatum'' gains entry and infects the root of the carrot. It is postulated that the sequence of enzymic production, in association with a slow growth rate, affects the plant infection response which causes the types of symptoms characteristic of this ''
Pythium ''Pythium'' is a genus of parasitic oomycetes. They were formerly classified as fungi. Most species are plant parasites, but ''Pythium insidiosum'' is an important pathogen of animals, causing pythiosis. The feet of the fungus gnat are frequen ...
'' species.


References


External links


USDA ARS Fungal Database
{{Taxonbar, from=Q7263902 Water mould plant pathogens and diseases Carrot diseases sulcatum