Mycobiota of Estonia

Seentel on looduses oluline osa - orgaanilise aine lagundajatena on nad asendamatud ökosüsteemide aineringes. Niiviisi osalevad seened ökosüsteemide sekundaarses produktsioonis, luues ühtlasi maakera loodusressursse. Inimese praktilises tegevuses on seentel tohutu tähtsus nii negatiivses kui ka positiivses tähenduses. Piisab, kui mõelda söödavatele, sealhulgas viljeldavatele seentele, mitmesuguseid keemilisi aineid produtseerivatele liikidele, ravimseentele, mürkseentele, mükooside tekitajatele, fütopatogeensetele seentele, hallitusseentele jpt. Seepärast väärivad seened igakülgset tundmaõppimist ning oma igapäevases tegevuses tuleb meil nendega tõsiselt arvestada. Eesti territoorium pakub seente leviku uurimise seisukohast laialdasemat huvi Euroopas tervikuna. Tänu Eesti looduslikele (botaanilistele, geograafilistele, geoloogilistele) iseärasustele on meie ala omapäraseks ristumiskohaks boreaalsete ja nemoraalsete, mõningal määral ka pontiliste seeneliikide areaalidele. Seetõttu on Eesti seenestik koosseisult mitmekesine ja liigirohke. Eesti seentest on kahe sajandi vältel kirjutatud hulgaliselt nii teaduslikke kui populaarteaduslikke töid. Viimastel aastakümnetel on ilmunud rida raamatuid mitmesuguste seenerühmade kohta, sealhulgas ka ülevaated meie parimate söögiseente perekondadest. Kõiki Eesti suurseeni käsitlev raamat “Seened” (koostaja K. Kalamees) ilmus juba 1966. aastal. Eesti pisiseeni laiemale üldsusele tutvustavat kirjandust on seevastu napilt, ometi on näiteks seente poolt põhjustatud taimehaiguste tundmine nii põllumajandus- kui metsamajanduspraktikas väga oluline. Eesti seente loend on meil küll ilmunud juba kahe raamatuna “Eesti seente koondnimestik” (Järva & E. Parmasto, 1980; Järva, I. Parmasto & Vaasma, 1998), kuid need mõlemad kujutavad endast seeneliikide kommentaarideta nimestikku koos viidetega vastavale kirjandusele (kuni aastani 1990). Käesolev raamat annab ülevaate umbes 4/5 Eestis kasvavatest seeneliikidest, püüdes seejuures neid lühidalt iseloomustada süstemaatiliselt, ökoloogiliselt, levikuliselt, bioloogiliselt ning kasu või kahju seisukohast inimesele. Lisaks sellele on raamatus iseloomustatud erinevaid seente kasvukohatüüpe Eestis, meie seente geograafiat, ökoloogiat, seenekaitset ning seente osa inimese elus. Paraku ei ole probleemide käsitlus raamatus siiski täielik, kuna Eestis ei ole veel mitmeid seenerühmi nimetatud küsimustega seonduvalt läbi uuritud. Nii on näiteks toitumisrühmade, kasvukohtade ja levilatüüpidega seotud üldistav analüüs meil seni tehtud vaid lehikseente osas. Käesolev raamat on teaduslik teatmeteos Eesti seentest. See ei ole ei määraja ega mükoloogia õpik ning seetõttu ei leia siin põhjalikku käsitlemist seente ehituse, bioloogia, paljunemise ja eluviisi probleemid. Neid küsimusi vaadeldakse üksikute seenerühmade juures vaid sedavõrd, kuivõrd nad osutuvad vajalikuks Eestist leitud seente iseloomustamisel. Mükoloogiline oskussõnastik hõlbustab raamatu kasutamist. Võimalikult täpselt on raamatus seeneliikide kõrval viidatud ka nende peremeesorganismidele koos ladinakeelsete nimetustega. Eesti kodumaistele puu- ja põõsaliikidele, levinumatele köögi- ja põlluviljadele, viljapuudele ja marjapõõsastele ning samuti kodu- ja metsloomadele on tekstis viidatud ainult eestikeelsete nimedega, vastavad ladinakeelsed nimed tuuakse eri nimestikuna raamatu lõpus. Raamatu töömahukast ning aegavõtvast kirjutamisest, koostamisest ja toimetamisest on osa võtnud palju kutselisi ja mitmeid harrastusmükolooge, samuti teistegi erialade esindajaid. Raamatu koostajana ja peatoimetajana avaldan siirast tänu kõigile autoritele ja kaastoimetajatele, ingliskeelsete tekstide tõlkijale M.Roosile ning keelelisele korrektorile M. Johansonile, CD versiooni tegijale ja kujundajale I. Kübarsepale. Raamatu failide esialgse töötlejana väärib kahtlemata tänu OÜ Eesti Loodusfoto. Oma käsikirjaliste materjalide kasutamise võimaldamise ning samuti kaastöö eest mitmete erinevate lõikude sisulisel täiendamisel ja parandamisel pälvivad lisaks neile tänu A. Jakobson, K. Jürgens, A. Kalamees, L. Kalamees, M. Laane, T. Randlane ja I. Saar. Käsikirja teksti trükkimise ja vormistamise eest väärib siirast tänu M. Vaasma. Eriline tänu kuulub posthuumselt raamatu illustraatorile kunstnik Georg Štšukinile, kelle sule ja pintsli alt on tulnud värvitahvlid ning mustvalged joonised.An investigation into Estonian mycobiota, including taxonomy, ecology, distribution and data on its resources, is presented in this book. A contemporary interpretation of the distribution of Estonian fungi between the kingdoms of Eucaryota and their systematic arrangement based on the principles of Hawksworth et al.(1995), as well as the morphology, anatomy, ecology, phenology, distribution, the profit or damage from the human point of view are considered in detail at the level of different taxonomic units from phyla to species. Trophic groups and sites, and the peculiarities of the geographical distribution of fungi in Estonia are analysed. Edible mushrooms, their resources and cultivation in Estonia, the nutritive value and ways of preservation, mycetism, mycotoxicoses and mycoses in man and domestic animals, poisonous fungi and medical uses of fungi, plant diseases caused by fungi, and dyeing of textile fabrics with fungal pigments are treated in separate chapters. Particular attention is devoted to the principles of fungus protection and the species included in the Red Data Book of Estonia and to those under state protection. The priority in the fungus investigations in Estonia belongs to Fischer and Hupel (1777). That is particularly evident in the works of Fischer (1778, 1784, 1791), Grindel (1803), Friebe (1805), Weinmann (1836), Dietrich (1856, 1859), Bucholtz (1904, 1916), Lepik (1930, 1940), Witkowsky (1934), Leisner (1937, 1938). First more concrete data of scientific significance are found in Dietrich’s and Bucholtz’s works which contain studies on the fungi of the Baltic Region of that time. From the period of 1925–1943 very important are the mycological and phytopathological investigations of prof. E. Lepik. The research centres were situated in Tartu. On E. Lepik’s initiative a number of amateur mycologists like N. Witkowsky, T. Leisner, A. Rühl and V. Pärtelpoeg joined in the work. In Estonia, the investigation into the systematic, ecology, distribution, pathology and coenology of fungi became more active in the 1950s, when studies were started by E. Parmasto, V. Lasting, P. Põldmaa, K. Kalamees. In the 1960s and later A. Raitviir, L. Järva, A.-L. Sõmermaa, M. Hanso, H. Karis, J. Sarv, K. Kask, B. Kullman, I. Parmasto, M. Vaasma, T. Normet, H. Lõiveke, P. Soobik joined in the research. The amateur mycologists H. Kelder, G. Shtshukin, V. Liiv and S. Veldre also took up training for mycological investigations. At present the research centre of Estonian mycology is the Institute of Zoology and Botany (department of mycology) by the Estonian Agricultural University. A new generation of mycologists, among them U. Kõljalg, K. Põldmaa have appeared. The traditional classical direction in Estonian mycological research is being replaced by a new one at the genetical and molecular level. Since 1777 almost 4000 species of fungi have been recorded in Estonia (cf. Järva, 1982; Parmasto, 1989). They have been treated in nearly 2000 books and articles (cf. Järva & E. Parmasto, 1980; Järva, I. Parmasto & Vaasma, 1998). Since 1950, 159 000 specimens of fungi have been collected for the fungus herbarium of the Institute of Zoology and Botany. The Estonian Mycological Society (until June 2000 the mycology section of the Estonian Naturalists' Society) has about 30 members. The composition of Estonian mycobiota is diverse, rich in species and resources, since Estonia lies in the temperate mixed forest zone of the northern hemisphere (Kalamees, 1995). In Estonia we find favourable growth conditions for both boreal coniferous forest fungus species and nemoral deciduous forest species. As the northern border of the distribution area of oak runs through South Finland, there are good growth conditions for practically all the fungus species connected with oak in Estonia. This is the reason why our mycobiota is considerably richer in comparison with that of the other northern countries. The development and character of Estonian mycobiota have been, to a great extent, influenced by the differences between the geological history of West ad East Estonia as well as the peculiarities of the soils, flora and climate in these regions. The differences in the base rock of North and South Estonia are equally important. West and North Estonia, including the islands of the Baltic Sea, considerably differ from South and East Estonia from the mycogeographical point of view as concerns, at least, mycorrhizal Agaricales. As regards the species composition of Agaricales, West and North Estonia are similar to Central and even to South Europe. The most important factors from the point of view of fungi in West and North Estonia are the abundance of broad-leaved tree species and calcareous soils on the Silurian and Ordovician limestone base rock. For these reasons Estonia seems to occupy an important position on the eastern and northern (north-eastern) borders of many nemoral fungus species spread in West Europe. Sometimes, however, the eastern (north-eastern) and northern borders of those species run in the close vicinity of the territory of Estonia, in Russia and South Finland, respectively. Owing to all the factors mentioned above the species composition of fungi in West and North Estonia is richer and more varied than that in East and South Estonia (which is also the case with the flora of higher plants). For instance, nearly 50 species of Estonian agarics grow only (or preferably) in West and North Estonia and on the islands (Kalamees & Lasting, 1973c). The mycobiota of Estonian forests is characterized by the domination of mycorrhizal fungi and litter saprobes (Kalamees, 1980a,c, 1982). There are few humus saprobes among forest fungi, their role is more important in forest type groups with a weak or missing litter horizon, such as alvar, dry boreo- nemoral, fresh boreo-nemoral, floodplain and paludified forests. The existence of wood saprobes and parasites, as well as leaf- and needle-debris saprobes is very characteristic of forest mycobiota. In the formation of the mycobiota of forest type groups the carbonate content of soils and their moisture regimes are of paramount importance. The mycobiota of Estonian meadows consists of humus saprobes, and lots of mycorrhizal fungi in parkland meadows and litter saprobes in denser stand groups (Kalamees, 1979c, 1980a, 1982). In parkland meadows quite frequent are also wood saprobes. The composition of the mycobiota of meadows is, to quite a great extent, influenced by human activities, mainly by grazing cattle and mowing. Pastures are, for instance, always rich in coprotrophs. The determining factor in the fungus composition of dry and fresh meadows is the carbonate content of the soils. The poverty of mycobiota in paludified meadows mainly accounts for excessive moisture. The mycobiota of Estonian mires basically consists of hygrophilous humus and moss saprobes (Kalamees, 1982). As concerns forested mires a few mycorrhizal fungi are also found. The main factor determining the character of mire mycobiota is the continuous excess moisture. The calcareousness of the soils does not exert significant effect on the mycobiota of mires. The mycobiota of Estonian boreal heath grasslands is very poor in species due to the extremely infavourable growth conditions (Kalamees, 1980a). The raw-humus nature of the forest litter horizon and high acidity do not create necessary conditions for the development of litter saprobes. The mycobiota of coastal dunes is poor in species but very peculiar in its species composition: psammophilous humus saprobes and xerophilous mycorrhizal fungi of willows and pines grow there. Sandy inland plains, which represent secondarily outcropping unfixed sands, are dry and therefore offer favourable growth conditions for only a few fungus species. Vegetation of outcrops in Estonia as well as halophilous coastal areas, off-shore bars, nitrophilous areas at the nesting places of birds, etc. are also very poor in fungus species (Kalamees, 1980a). A number of water fungi grow in Estonia. They are found on plant remnants deposited on the bottom of water-bodies, on stalks of live plants and on other organic substrate in water. They mainly belong to Hyphomycetes (A. Kalamees, 1989). Macrofungi cannot grow in water, but favourable conditions for the development of many Helotiales are created in reed-beds and other groups of water plants as a result of the accumulation of decaying plant remnants after the flood has sunk. The mycobiota of ruderal and cultivated vegetation is highly varied and peculiar (Kalamees, 1981). The basic factor determining its composition is the humus content in the soil. Mainly humus saprobes grow on these sites, but coprotrophs are also often found. Lots of mycorrhizal fungi grow in parks. Estonian mycobiota is rich in edible fungi being represented by almost 400 species. The general resources of Estonian fungi reach to 36.5 thousand tons (Kalamees & Vaasma, 1980). Among forest types with stands ready for felling the following types can be undoubtedly considered the most productive: Vaccinium uliginosum pine type with 229 kg per hectare, Cladonia pine type with approximately 215 kg per hectare and Calluna pine type with approximately 239 kg per hectare (Kalamees & Vaasma, 1980; Kalamees & Silver, 1988, 1993). According to the latest data it is the young Cladonia type (25 years old) that with 569 kg per hectare exceeds manyfold the fungus yields of any other Estonian forest site. The most productive forests, as concerns the fungus yield, lie in North, South-East and South-West Estonia, and on the island of Saaremaa. As concerns fungus species, the following can be considered to be the most productive: Lactarius rufus with 495 kg per hectare, Suillus bovinus with 165 kg per hectare, Suillus variegatus with 129 kg per hectare and Russula decolorans with 94 kg per hectare (Kalamees & Silver, 1988). Among the 200 species of poisonous macrofungi have been recorded in Estonia, there are three deadly poisonous ones: Amanita virosa, A. phalloides and Inocybe erubescens. Mycetism has been rare in Estonia. Amanita virosa was the reason of four poisonings causing death during the period of 1935–1998 (see Lepik, 1935a; Witkovsky, 1935). Some quite serious poisonings were caused by Inocybe erubescens, Gyromitra esculenta, Cortinarius sp. (subg. Phlegmacium, sect. Xanthophylli), Paxillus involutus and Phaeolepiota aurea in recent years. Relatively many cases of mycotoxicoses in cattle recorded in Estonia during the last half a century were caused by spoilt feed. Dermatomycoses and candidiasis quite wide-spread mycoses are in people and domestic animals and aspergillose and trichohytosis in domestic animals and poultry. In Estonia there are about 150 species of macrofungi belonging to medical fungi, 3/4 of them for their antibiotic qualities. In fact, only 2 species have been used for this purpose. Claviceps purpurea is the only pharmacological fungus medicine used. In folk medicine Inonotus obliquus is used in the cure of cancer. There are about half a thousand fungus species in Estonia which pigments could be used for dyeing textile fabrics. However, as a matter of fact, they have never been used for this purpose. The basic principle of the fungus protection in Estonia consists in the protection of fungus habitats (Kalamees, 1988). As a result, we can protect successfully both the fungus resources and separate species requiring protection. The Red Data Book of Estonia contains 91 fungus species, the list of fungi under protection contains 30 species (Kalamees & Vaaasma, 1998).Käsikirja valmimist rahastas Eesti Teadusfond ja Eesti Haridusministeerium

Inorganic mercury accumulation in brain following waterborne exposure elicits a deficit on the number of brain cells and impairs swimming behavior in fish (white seabream - Diplodus sargus)

The current study contributes to fill the knowledge gap on the neurotoxicity of inorganic mercury (iHg) in fish through the implementation of a combined evaluation of brain morphometric alterations (volume and total number of neurons plus glial cells in specific regions of the brain) and swimming behavior (endpoints related with the motor activity and mood/anxiety-like status). White seabream (Diplodus sargus) was exposed to realistic levels of iHg in water (2 mu g L-1) during 7 (E7) and 14 days (E14). After that, fish were allowed to recover for 28 days (PE28) in order to evaluate brain regeneration and reversibility of behavioral syndromes. A significant reduction in the number of cells in hypothalamus, optic tectum and cerebellum was found at E7, accompanied by relevant changes on swimming behavior. Moreover, the decrease in the number of neurons and glia in the molecular layer of the cerebellum was followed by a contraction of its volume. This is the first time that a deficit on the number of cells is reported in fish brain after iHg exposure. Interestingly, a recovery of hypothalamus and cerebellum occurred at E14, as evidenced by the identical number of cells found in exposed and control fish, and volume of cerebellum, which might be associated with an adaptive phenomenon. After 28 days post-exposure, the optic tectum continued to show a decrease in the number of cells, pointing out a higher vulnerability of this region. These morphometric alterations coincided with numerous changes on swimming behavior, related both with fish motor function and mood/anxiety-like status. Overall, current data pointed out the iHg potential to induce brain morphometric alterations, emphasizing a long-lasting neurobehavioral hazard.“Fundação para a Ciência e a Tecnologia” (FCT). This work has been supported by the Research project financed by FCT PTDC/AAG-REC/2488/2012 (NEUTOXMER - Neurotoxicity of mercury in fish and association with morphofunctional brain alterations and behavior shifts), as well as by the Centre for Environmental and Marine Studies (CESAM)