There has not been complete agreement as to what is meant by ectendotrophic mycorrhizae, and there is a wide variety of opinion among authors on mycorrhizal terminology. In this paper ectendotrophic mycorrhizae are defined to be short roots with Hartig net and intracellular hyphae in the cortex. A mantle and digestion of intracellular hyphae may be found but are not necessary. In the study of Mikola (1965) ectendotrophic mycorrhiza was found to be common in Scots pine (Pinus sylvestris L.) seedlings in Finnish nurseries. The mycorrhizae had always similar structure and the mycelium isolated from the seedlings (E-strains) was similar. The aim of this study was to find out what kind of ectendotrophic mycorrhizae exist in forests and nurseries outside Finland, what kind of mycorrhizae do the E-strains isolated from Scots pine form with other tree species, and are these associations symbiotic.
Only one type of ectendotrophic mycorrhiza was found on the 600 short roots collected from the continents of Europa and America. The type was similar to the one described by Mikola: the mycelium is coarse and forms a strong Hartig net, and intracellular infection is heavy. Evidence is convincing that this structure was formed by the same fungus species. The species is unidentified. Mycorrhizae synthesized by E-strain with six spruce species, fir, hemloch and Douglas fir were all ectotrophic.
The E-type ectendotrophic mycorrhizae proved to be a balanced symbiosis. The seedlings of 13 tree species inoculated with the E-strain grew in the experiment better than the controls. The observation that ectendotrophic mycorrhizae dominates in the nurseries but is seldom found in forests, and then only in seedlings growing in the forest, was confirmed in the study. In synthesis experiments E-strain formed either ecto- or ectendotrophic mycorrhiza depending on the tree species.
The differences between different types of mycorrhiza; endomycorrhiza, ectomycorrhiza and ectendomycorrhiza, and the use of the terms have been variable in the earlier research. Studied of mycorrhiza in Scots pine (Pinus sylvestris L.) and Norway spruce (Picea abies (L.) H. Karst.) seedlings may suggest that the conditions affect which kind of mycorrhiza develops in the seedlings. This study is aimed mainly at finding out whether the difference of ectotrophic and ectendotrophic mycorrhizae depends on fungal symbionts or envirionmental conditions. Furthermore, the occurrence of ectendotrophic mycorrhiza in Finland under various conditions was studied, and experiments on the physiology and ecology of the mycorrhiza and the fungal partner were conducted.
The ectendotrophic mycorrhiza as described in this paper has proved to be very common on Scots pine in Finnish nurseries, but it was not found in Norway spruce seedlings. The results did not support the hypothesis presented in some earlier studies that ectendotrophic mycorrhiza is more parasitic than the other mycorrhizal fungi. The nursery survey showed that no correlation existed between the size and vigour of the seedlings and the presence of ectendotrophic mycorrhiza. Furthermore, greenhouse-grown seedlings with and without the fungus grew equally well. The type of mycorrhiza was, however, almost exclusively confined to young (1–3-years-old) seedlings and to nursery soils. The experiments indicates also that ectendomycorrhizal fungus has a very wide ecological amplitude in regard to light intensity, soil fertility, acidity, and humus content. It has, however, a weak competitive ability in natural forest soils against the indigenous fungal population. When the seedlings were transplanted from the nursery to forest soil, their mycorrhizal population was largely changed.
Prescribed burning is a common silvicultural practice in northern Europe, intended to destroy the slash and ground vegetation and to reduce the thickness of the raw humus layer prior reforestation. The purpose of the experiments was to study whether there are any differences in the commencement and early development of mycorrhizal infection between burned and unburned areas. A clear-cutting area was burned on May 1961. The soil was rocky moraine, the forest type was Vaccinium type. Two weeks after burning Scots pine (Pinus sylvestris L.) was sown in patches.
According to the results, mycorrhizal infection took place on the unburned area earlier than on the burned. The difference was relatively small, perhaps 1–2 weeks. Although burning kills mycorrhizal fungi, it did not cause serious harm to the seedlings, on the contrary, the favourable influence of burning was more distinct. The high temperatures caused by the fire are restricted in the soil in a prescribed burning only a few centimetres deep. Although the mycorrhizal fungi are concentrated in a very thin surface layer of the soil, some mycorrhizae are situated deeper, and from there the fungi are able to infect roots and spread back to the surface layer. The fire also rises the pH of the soil, which can be harmful for mycorrhizal fungi. Even this effect, however, is limited to a thin surface layer.
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Mycorrhizal association is a characteristic feature of the trees of the northern coniferous forests. The purpose of the present study was to determine what influence some fungicides and herbicides regularly used in Finnish nurseries have on formation and development mycorrhizal in Scots pine (Pinus sylvestris L.) and Norway spruce (Picea abies (L.) H. Karst.) seedlings. The results are based mainly on field experiments in nurseries. First the initiation of mycorrhiza was described in untreated seedlings.
In the first growing season mycorrhizal infection commences fairly late even under normal conditions, i.e. 6–7 weeks after seeding and 3–4 weeks after the formation of the first short roots. Soil disinfectants are commonly used in nurseries before seeding, and they are supposed to evaporate or disintegrate in a few days or 1–2 weeks. In pure culture experiments mycorrhizal fungi proved several times more sensitive than parasitic and indifferent soil moulds to herbicides and fungicides, but in field experiments the delay of mycorrhizal infection caused by them does not seem to harm the seedlings. In the second summer differences of mycorrhizal relations between treated and control plots disappeared. Accordingly, the influence of biocides on mycorrhizae, when applied in the customary concentrations, does not extend beyond the first growing season.
Methyl bromide and SMDC retarded mycorrhiza formation distinctly, while formaldehyde and allyl alcohol had no effect. Apart from not retarding mycorrhizae, formaldehyde and allyl alcohol promoted seedling growth and favoured Trichoderma viride in the soil. Trichoderma is known to be antagonistic to many fungi.
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Draining transforms root systems of trees growing in peatlands towards the ones growing on mineral soil. However, even after efficient draining the root systems differ from the root systems of trees growing on mineral soil. This investigation concentrates on root systems of forests of similar mire types growing in similar draining conditions but having different tree species compositions. The peatland, situated in Pieksämäki in Southern Finland, was drained in 1937. Sample plots, measured in 1956, consisted of mixed forest of Scots pine (Pinus sylvestris L.), Norway spruce (Picea abies L. Karst.) and birch (Betula sp.) in different compositions, and were in natural condition.
The sedge pine bog studied in this investigation was shown to have larger total amount of roots and mycorrhiza than in previously studied dwarf shrub pine bogs. This reflects better growth conditions of the better site. The depth of root system was, however, similar. Root systems of birch were deeper than those of the coniferous tree species. Differences between Scots pine and Norway spruce were small. Corresponding differences between the species were found in the density and total number of mycorrhizas. The abundance of mycorrhizas in the roots of birch increased in deeper layers of peat, but decreased especially in spruce roots. In earlier studies the abundance of mycorrhizas decreased in the roots growing in deeper layers in pure Scots pine stands, but no such variation was seen in this study. The result suggest that the deep root system of birch may affect also the root systems of the coniferous trees. On the other hand, birch roots can have advantage over the coniferous trees.
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The root system of a Scots pine (Pinus sylvestris L.) growing on a peatland is restricted, according to earlier studies, on the top layers of the peat above the groundwater level. Drainage of the peatland affects growth of the root system. This investigation aims at studying the root systems on the point of view of draining of peatlands. The structure and distribution, and the growth of mycorrhiza in Scots pine roots in pine swamps varying from natural state to well drained state is studied.
The study shows that Scots pine on pine swamps has more extensive root system than has earlier assumed, it is common to find 1,000 m of roots in one cubic meter in a healthy stand. The trees reach this density of roots early on. In a drained peatland, the total root length is markedly higher than in a similar stand in natural state. The root systems proved to be very shallow. Even in a well-drained site the roots did not grow deeper than 20 cm. 70% of all roots were found in the upper 5 cm layer of peat, and 90% in the upper 10 cm layer. Root systems were deeper in drained peatlands, but the difference was small. In a site in natural state the average depth of the roots was 4 cm, and in a drained site 5 cm. About 85% of the roots were under 1 mm of diameter. Short roots were found only in the fine roots. Draining increases strongly the number of short roots. Mycorrhizas of the types A, B, C and D as well as pseudomychorrizas were found in the pine roots.
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While the most common type of mycorrhizae is endomycorrhizae, ectomycorrhizae dominate in the case of coniferous trees. Pine, in particular, has a strong association with mycorrhizae. Mycorrhizae enable trees to take up water and nutrients much more efficiently than the roots themselves. The fungus, in return, obtain carbohydrates and is able to grow and fruit. Mycorrhizal fungi are probably numbered in their thousands but so far few are known. Knowledge about their physiology, in particular, is lacking and studies dealing with their isolation and inoculation, which may be commercially valuable, remain unpublished. A new challenge for mycorrhiza research is the effects of air pollution. Forest suffering from extensive air pollution have few mycorrhizal fungi., infection is weak and the number of root deformations is high. As good mycorrhizae are important to tree health, there is a particular need to intensify mycorrhiza research.
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While the most common type of mycorrhizae is endomycorrhizae, ectomycorrhizae dominate in the case of coniferous trees. Pine, in particular, has a strong association with mycorrhizae. Mycorrhizae enable trees to take up water and nutrients much more efficiently than the roots themselves. The fungus, in return, obtain carbohydrates and is able to grow and fruit. Mycorrhizal fungi are probably numbered in their thousands but so far few are known. Knowledge about their physiology, in particular, is lacking and studies dealing with their isolation and inoculation, which may be commercially valuable, remain unpublished. A new challenge for mycorrhiza research is the effects of air pollution. Forest suffering from extensive air pollution have few mycorrhizal fungi., infection is weak and the number of root deformations is high. As good mycorrhizae are important to tree health, there is a particular need to intensify mycorrhiza research.
The PDF includes an abstract in English.
The fungal symbiont of ectendomycorrhizae is an ascomycete Wilcoxina (Tricharina) mikolae Yang & Korf. It forms ectendomycorrhizae with Pinus and Larix and ectomycorrhizae with Abies, Picea, Pseudotsuga and Tsuga. It is common in forest nurseries around the world. After transplanting the seedlings into natural forest soil, indegenous fungi rapidly replace Wilcoxina. Inoculation of nursery soil with Wilcoxina is recommended if soil has been sterilized or for other reasons mycorrhizal fungi are absent.
The PDF includes a summary in Finnish.
The fungal symbiont of ectendomycorrhizae is an ascomycete Wilcoxina (Tricharina) mikolae Yang & Korf. It forms ectendomycorrhizae with Pinus and Larix and ectomycorrhizae with Abies, Picea, Pseudotsuga and Tsuga. It is common in forest nurseries around the world. After transplanting the seedlings into natural forest soil, indegenous fungi rapidly replace Wilcoxina. Inoculation of nursery soil with Wilcoxina is recommended if soil has been sterilized or for other reasons mycorrhizal fungi are absent.
The PDF includes a summary in Finnish.
The host range of Paxillus involulutus includes a wide range of species. These mycorrhizae can be identified in the field by their appearance. A positive correlation was found between the numbers of mycorrhizae and sporophores formed by the species. It is concluded that Paxillus involutus does not form sporophores when growing by a saprophytic mode of nutrition. In the presence of trees, the species fruits to varying extents: in poor closed stands hardly at all and in fertile stands profusely. After partial cutting, soil scarification, draining and application of nitrogen, its fruiting increases markedly. Consequently, growth of Paxillus involutus in raw humus is arrested primarily due to deficiency of nitrogen.
In pure culture the amount of submerged mycelium on agar is very limited, but the aerial mycelium profuse. In the latter, sclerotia are also formed. The pH and temperature requirements may vary between individual strains. The species is also able to utilize starch. Nitrogen is utilized in the form of both ammonium and nitrate, and organic nitrogen sources.
Paxillus involutus forms a balanced symbiosis, even when the host is relatively weak and the fungus relatively virulent. It survives rather well in Scots pine seedlings planted in various sites; moreover, the initial development of these seedlings is better than that of nonmycorrhizal seedlings.
This study emphasizes the need for thorough investigations concerning whether mycorrhizal fungi are capable of fruiting when subsisting by a saprophytic mode of nutrition. In pure culture experiments several strains should be used. Semi-aseptic synthesis is sometimes surprisingly rapid, its major handicap being the limited number of fungal symbionts that can be successfully inoculated. In both this and aseptic synthesis mycorrhizal associations can be formed whose existence in nature is questionable.