Tafrina

E. Fries in 1815 described a genus called Taphria , which is formed from ancient Greek. τάφρη "moat" ^[1] . In later works by Vries and other authors, the name Taphrina is used .

The asexual reproduction stage ( anamorph ), or tafrin yeast, was described as an independent genus in 1990 ^{[* 1]} . They are given the name Lalaria , given on the eponymous pebble beach ( Greek Λαλρια ) on the Greek island of Skiathos . Lalaria Beach is known for its smooth stones of the same size. The view of the yeast cells under a microscope resembles this pebble ^[2] .

Taffrin plum. Figure 1892
A) affected plum fruit (“pocket”)
B) microscopy of the cut surface of the “pocket”: m - mycelium of the fungus; e - plant cells; s - sporiferous layer; c - fetal cuticle
C) Aski with a higher magnification: a - developing; b - with matured ascospores ; st - basal cell
D) disputes after release from ask

Fruit bodies and structures of asexual sporulation (conidioma) are not formed.

Mycelium develops in the tissues of infected plants. Mycelium is dikaryotic, that is, consists of binuclear cells. Cell walls are bilayer. In some species, well-developed infectious structures are formed, penetrating into the cells of the host plant, haustoria . During sporulation, the mycelium forms a layer under the cuticle (outer protective film) of the plant, the cells of this layer germinate with the spore-bearing organs — asci (bags) ^[3] . Partitions between cells of the mycelium ( septa ) are characterized by the presence of numerous micropores; the accompanying organelles found in most of the fungi near the septal pores are absent from taffrins ^[4] .

Aski eutunikatnye (with a two-layer shell), colorless, short, most often cylindrical or club-shaped in shape with a truncated or rounded top. Aski form a dense layer, similar to the hymenia and often called, but unlike the true hymenia of the Ascomycetes, it never contains sterile elements. After the bags are formed, the cuticle of the plant bursts and the marsupial layer is exposed. The marsupial layer is located on the surface of the affected parts of the plant, often on the underside of the leaves, it has the appearance of a powdery or wax-like patina of gray, whitish or golden color. In some species (for example, Taphrina potentillae ) bags do not form a continuous layer, but are arranged in separate sections. Eight ascospores usually mature in Askah , less often four ^[5] .

At the base of the bags, many species have basal , or subnumber, cells (“leg cells”). They are less than bags in length, and differ in width in different types — they can be wider or narrower than bags. The basal cells are separated from the bags with septum without pores ^[6] .

Ascospores, called in these fungi also endospores , are unicellular, colorless, smooth, spherical, ellipsoidal, or rod-shaped. They germinate, forming smaller secondary blastospores , or conidia . Blastospores multiply by budding and form pink or red yeast colonies on nutrient media, similar to some basidiomycete yeast ^[7] . Secondary disputes in some authors are called sporidy or oidy ^[8] .

The nuclear life cycle - haploid - dikaryotic , dominated by the dikaryotic phase ^[9] .

Haploid ascospores germinate with blastospores; in some species, ascospores germinate before release from bags. Blastospores give rise to an anamorphic haploid yeast phase consisting of budding cells that are saprotrophic . Tafrin yeast is described in the composition of the anamorphic genus Lalaria , introduced in 1990 ^[10] . The haploid phase ends with a copulation (fusion) of cells and the formation of dikaryon. In some species, dikarization can occur without mating cells, by dividing the nucleus. In artificial breeding, dikaryotic mycelium usually does not develop, but the appearance of particular thick-walled cells is noted. It is believed that these cells are able to survive under adverse conditions, that is, they perform the function of chlamydospores in nature. Dikariotic mycelium, unlike most other ascomycetes (except saccharomycetes ), feeds on its own, and not at the expense of the haploid phase. Dikariofaza leads exclusively parasitic (biotrophic) lifestyle, developing between the cells of an infected plant ^[11] .

For the genus is characterized by a specific method of forming bags, called Taphrina- type . Under the cuticle of the plant, a layer of cells is formed, called ascogenic cells , which are fragments of undifferentiated hyphae. They merge nuclei (karyogamy), diploid cells increase in size and their nuclei share mitosis . Simultaneously with mitosis, the ascogenic cell differentiates into the basal and pre-aspens ( procreate ) cells, each of which receives one daughter diploid nucleus. Then, meiosis takes place in the pre-husky cell, and a short 8- or 4-spore bag is formed ^[12] , after sporulation, the basal cell degenerates ^[11] .

Basal cells are not formed in all species; also bags with their own shells may form inside the ascogenic cells, the walls of the latter then burst out. The formation of ascospores also has some features that distinguish taffrin from other ascomycetes. Each of the haploid nuclei with parts of the cytoplasm is surrounded by a plasma membrane that is drawn into the inside of the bag from its wall, then the membrane forms a spore wall ; plasma membrane-independent membrane sac is absent. After the spore has matured, the top of the bag is opened with a simple slit, and the spores are released passively or thrown away with force ^[11] .

The genus is distributed mainly in the temperate zone of the Northern Hemisphere , but it is noted that the species composition is poorly studied in the tropics and subtropics . To the north, representatives of the genus penetrate up to the border of the range of their host plants. In the southern hemisphere , the species Taphrina entomospora , infecting Nothofagus pumilio on Tierra del Fuego , is known; several other species are widely distributed in South Africa ^[13] .

Typhrina species are obligate parasites of higher plants, which determines the importance of many of them as pathogens of plant diseases used in the human economy. Diseases of garden trees caused by taffrins, such as witch brooms , leaf curliness and fruit damage ( pockets or blown fruits ), can cause significant damage. Such diseases are found in stone fruit trees (representatives of the Plum genus) - homemade plums , ordinary cherries , peaches , almonds, and others; pear leaves may suffer from leaf curl ^[14] .

Disease curliness of leaves and "witch brooms" leads to a decrease in yield, weakening of trees and infection by secondary infections , can lead to the death of plants, and with the defeat of the fruit, a direct loss of part of the crop occurs. Wild trees and shrubs that are of economic importance - the turn , the bird cherry - are also affected by these diseases ^[14] .

In addition to garden trees, forest and park trees such as birch , maple , alder , oak , elm , and poplar can suffer from diseases of the leaves, flowers, and fruits. The appearance of witch bristles and leaf diseases in urban plantings leads to a decrease in their ornamentation, forest trees from taffrin diseases slow down growth and become more sensitive to frost. Infection of forest species often occurs through damage by insects ^[14] .

Symptoms of diseases caused by

According to morphological features, the types of taffrins often do not clearly differ from each other ^[15] , therefore the nature of the damage and the taxonomic group of host plants are important for taxonomy of the genus and the identification of species. Damage to tafrins can lead to profound disturbances in the physiology of the plant , the appearance of various deformities (deformities, hypertrophy of organs, underdevelopment of seeds, pathological growths). These disorders are caused by the ability of fungi to produce substances that have hormonal activity or stimulate the synthesis of hormones by plants. Tufrins synthesize heteroauxin and other substances of the auxin class, as well as cytokinins . In the 1960s, Japanese scientists isolated an active substance that was different from heteroauxin; the name tafrinin was suggested for it ^[16] .

The active substances of the fungus stimulate the growth and division of plant cells and inhibit the processes of cell differentiation.

In cases of mycelium development under the cuticle, deformations usually do not appear, the lesion is limited to the appearance of spots of various sizes and shapes on the leaves , stems , and wyai ferns. Spotting occurs as a result of tissue necrosis , which may be accompanied by cell hypertrophy . In the latter case, a thickening appears on the upper side of the affected leaf, and on the lower side there are depressions covered with a mushroom pouch ^[16] .

When the mycelium develops intercellularly, the leaves remain morphologically underdeveloped, but at the same time they grow and deform, acquire an uncharacteristic color, the chlorophyll in the affected leaves is gradually destroyed. The veins of infected leaves remain shortened, and the plate grows about 2 times compared with healthy leaves, thickens and becomes fragile. The stomata lose their ability to close; as a result, the normal breathing and the water balance of the plant are disturbed. This symptom is known as "curly". Affected leaves die off and fall off, which can greatly weaken the plant ^[17] .

The defeat of the flowers in plants of the Rosaceae family leads to the appearance of terry , pathological growth of receptacle , hypertrophy and underdevelopment of reproductive organs . In fruits, the pericarp grows strongly, their size can increase several times, and instead of a seed, a cavity is formed, such a lesion is called “exaggerated fruit” or “pockets”. In willow and birch , the scales of earrings grow and become deformed, and the fruit is bloated. The affected fruits and flowers are covered with a spore-borne bloom, and then fall off ^{[18] [16]} .

When infected, the fungus can stimulate the growth of axillary buds of lateral shoots, which leads to the appearance of “witch brooms,” sometimes reaching a diameter of 3 meters. Witch brooms consist of dense clusters of short and delicate shoots, often heavily branched. Such shoots almost never bear fruit, the leaves on them are small, irregular in shape, rapidly falling ^[19] .

Other symptoms include the curvature and thickening of the stems, the proliferation of the parenchyma of the cortex and change in its color, damage to the vascular system ^{[16] [14]} .

Ways to fight

Mycelium of taffrin fungi is always in the tissue of infected plants, so it is difficult to cure a sick tree. Damaged branches are pruned and destroyed; they also destroy the fallen fruits with their “pockets”, flowers and leaves. Wounds after pruning are covered with garden pitch, diseased trees are treated with fungicides .

To combat curliness, “pockets” in early spring and autumn, trees are sprayed with 1–2 percent Bordeaux liquid or 0.75 percent copper sulphate , in the summer also Bordeaux liquid is used in 1 percent concentration, 0.3—0.4 percent copper oxychloride or sulfur preparations (tiovit) ^[14] .

In the modern systematics adopted in the Dictionary of Einsworth and Bisbee Mushrooms , the Taffrin family ( Taphrinaceae Gäum. & CW Dodge 1928 ) with the only Tafrin species ( Taphrina Fr. 1815 ) is included in the order Taffrin or Taffrin ( Taphrinales Gäum. & CW Dodge 1928). ), containing another family - Protomytsieva ( Protomycetaceae ). The Taffrin Order is a monotype class of Taphrinomycetes ( Taphrinomycetes OE Erikss. & Winka 1997 ), which, together with three other classes, is a subdivision of the Taphrinomycotina OE Erikss. & Winka 1997 Department of Ascomycetes ( Ascomycota Caval.-Sm. 1998 ).

In the old systems, the family was called Exoascaceae G. Winter 1884 , and the order - Exoascales ^[20] - by one of the genera ( Exoascus Fuckel 1860 ), later combined with the Tafrin family. In 1992, the Committee on Mushrooms and Lichens of the International Botanical Congress banned the name Taphrinaceae instead of Exoascaceae ^[21] .

Many authors divided the family into several genera. So, K. Fukel included in the genus Taphrina s. str. species that have one-year mycelium and cause blotchiness and slight leaf deformations, while species with a perennial mycelium that cause deeper lesions are attributed to the genus Exoascus . The genus Exoascus was recognized by R. Sadebeck and A. A. Yachevsky ^[22] . Species in which the aski are grouped into separate bundles and do not form a continuous sporiferous layer were attributed to the genus Magnusiella ^[23] . N. Century Sorokin , the founder of the study of taffrin in Russia, identified in the family of three genera - Taphrina (species with mycelium, developing under the cuticle), Exoascus (species with intercellular mycelium) and Ascomyces (no mycelium) ^[24] .

There have also been attempts to isolate subgenera in the Taffrin genus, for example, on the basis of the specialization of species for various host plants. K. Gisenhagen in 1901 identified the subgenera Taphrinopsis (parasitic species on ferns), Eutaphrina (on willow, birch and elm) and Euexoascus (on pink) ^[22] .

Subsequently, such fragmentation of the family and genus were considered artificial and rejected.

The position of the taffrin family (or Exoascaceae ) in the mushroom system has long been uncertain or controversial. In the 20th century, the Ascomycetes, which form fruit bodies, were assigned to the Euascomycetes group ( Euascomycetes ), which was usually considered as a subclass, and more primitive mushrooms were assigned to the Protoascomycetes (“protosoamed”) or Gymnoasci (“hollowed”) subclass ^[25] , and later - hemiacecomycetes ( Hemiascomycetidae ) ^[26] . In addition to taffrin, this subclass contained yeasts of the order Protascales or Endomycetales , now referred to the class of saccharomycetes ( Saccharomycetes ) and some other groups of fungi. Such a merging of all “voice-voiced” into one taxon was considered formal, it was noted that the presence of tamarino mycelium, eutunicular bags and the active ejection of the spores brings them closer to the Euaisceomycetes. It was assumed that tafrin phylogenetically associated with discomycetes (in the modern systematics, the subdivision Pezizomycotina , which includes the classes Pezizomycetes and Leotiomycetes ), and the simplification of morphological signs was the result of adaptation to parasitism ^{[27] [28]} . A supporter of the origin of tafrinovs from discomycetes was, for example, E. A. Goymann ; DK Zerov rejected this assumption, pointing to the simpler structure of tafrinov asks ^[29] .

There were also points of view on tafrin ones as the most primitive group from which euasicomycetes originated ^[30] or as a specialized branch of protosoism that deviated from the original forms in connection with parasitism ^[28] .

Another popular notion in the 20th century is that taffrin ones are closer to basidiomycetes than to marsupials. Adherents of this hypothesis considered the bag of a taffrin mushroom as a degenerated basidium (for example, G. Lovag ). D.Sevil believed that taffrin are the ancestors of basidiomycetes, and on the other hand, associated them with mukorovymi fungi ( zygomycetes ). A. Tleler in 1988 determined the family of taffrin to the class described by him Protobasidiomycetes , which also included the family of sugar moths ( Saccharomycetaceae ). Y.A. Arks , on the contrary, all marsupial, basidiomycete and deuteromycete yeast and parasitic fungi with yeast stages were classified as ascomycetes Endomycetes ^[30] .

In the 1990s – 2000s, numerous molecular phylogenetic studies were carried out, showing that tafrin and some other groups of fungi are a branch that has moved away from the rest of the Ascomycetes in the early stages of their evolution, that is, they constitute the basal group of Ascomycetes. These groups of fungi are now united in the Taphrinomycotina subdivision. The Sister group of the subdivision is Saccharomycetes (subdivision Saccharomycotina ) ^[30] .

Tuffrin yeast was first described as an independent genus Saprotaphrina in 1962 by the Italian authors O. Verona (O. Verona) and A. Rambelli (Rambelli, Antonella). The Saprotaphrina taxa, however, were not really made public ( nom. Inval. ), Nor was it strictly proved that the objects studied were indeed asexual stage of taffrin species. In 1990, the anamorph was re-described by R.T. Moore (Moore, Royall T.) under the name Lalaria . Moore described 23 species, and in 2004 a group of Portuguese researchers described 5 more species, for 4 of which the mycelial stage was not detected. In this regard, the diagnosis of the genus Lalaria was revised, it was stated that the teleomorph, if it exists , belongs to the genus Taphrina ^[31] .

Types

The genus contains about 100 species ^[22] (according to the data of the Dictionary of Ainsworth and Bisbee ^[32] - 95). The Index Fungorum database contains 173 species and infravidous names of the genus Taphrina (including synonyms ).

Some species

• Taphrina betulina Rostr. 1883 parasitizes various types of birch . Earlier, a number of researchers stood out as an independent species of Taphrina turgida ; according to modern concepts, it is probably the earliest stage of development of Taphrina betulina . The disease begins with the appearance of yellow spots on the leaves, then the mycelium affects the shoots, which leads to the appearance of the “ witch bristles ”. Trees with a strong defeat "witch brooms" slow growth, produce fewer seeds. Taphrina betulina is found in the northern and mountainous regions of Eurasia and in Greenland. Mycelium in this species wintering ^{[33] [34]} .
• Taphrina bullata ( Berk. & Broome ) Tul. 1866 - Tufrin blistering causes leaf curliness in pears and related species of the Pink family. Affected plants are weakened by a decrease in the intensity of photosynthesis . Taffrine blister is found in Europe, the Caucasus and the Far East, is widespread in North America. Mycelium one-year ^[35] .
• Taphrina caerulescens ( Desm. & Mont. ) Tul. 1866 parasitizes on oak , causes spotting and deformation of young leaves. It has an annual mycelium and usually does not cause significant damage to plantations, but with repeated infection sometimes leads to the death of individual trees. Встречается в регионах с умеренным и субтропическим климатом Северного полушария ^[36] .
• Taphrina cerasi ( Fuckel ) Sadeb. 1890 вызывает «ведьмины мётлы» у вишни , а близкий вид Taphrina wiesneri ( Ráthay ) Mix 1954 — Тафрина Визнера , или малая — является возбудителем курчавости листьев. Эти два вида встречаются в Евразии, Северной Америке, Южной Африке и Австралии; имеют многолетний мицелий ^[37] .
• Taphrina deformans (Berk.) Tul. 1866 — Тафрина деформирующая — один из наиболее изученных видов, возбудитель курчавости листьев персика и миндаля . Имеет космополитическое распространение и может наносить заметный ущерб садоводству. Мицелий многолетний. Гриб, поражающий миндаль, некоторые авторы считают самостоятельным видом — Тафрина миндаля ( Taphrina amygdali ) ^[38] .

• Taphrina padi ( Jacz. ) Mix 1947 поражает плоды черёмухи обыкновенной и близких видов из рода Слива , вызывая появление « кармашков ». Широко распространена в Евразии, в Западной Сибири иногда вредит, полностью уничтожая завязи черёмухи. Имеет многолетний мицелий ^[39] .
• Taphrina populina ( Fr. ) Fr. 1832 — Тафрина золотистая — типовой вид рода. Вызывает заболевание листьев тополя («пузырчатку»). Листья при этом заболевании покрываются пузыревидными вздутиями, а затем отмирают. Тафрина золотистая может вредить городским насаждениям, снижая их декоративность. Мицелий однолетний ^[40] .
• Taphrina pruni Tul. 1866 — Тафрина сливовая , как и деформирующая, достаточно хорошо изучена. Она заражает многие виды сливы , относящиеся к подродам Слива и Вишня . Этот вид способен вызывать повреждения плодов («кармашки»), листьев (деформацию) и ветвей («ведьмины мётлы»). Может наносить ущерб, уничтожая в отдельные годы до 15—20 % урожая только в результате непосредственного поражения плодов, также ослабляет деревья, создавая благоприятные условия для вторичных инфекций. Мицелий многолетний. Встречается во многих регионах мира, как в Северном, так и в Южном полушариях, но не повсеместно. Ранее выделялся близкий вид Тафрина терносливы , который в современной таксономии не является признанным. Тафрину терносливы считали возбудителем «ведьминых мётел», а сливовую — «дутых плодов». ^[41] .

comments

1. ↑ В микологии допускалось описание отдельных стадий жизненного цикла как самостоятельных таксонов. Это правило отменено XVIII Международным ботаническим конгрессом в июле 2011 года.

ссылки на литературу

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