Marja Roitto

SEVERE DEFOLIATION OF SCOTS PINE REDUCES REPRODUCTIVE INVESTMENT BY ECTOMYCORRHIZAL SYMBIONTS

Reduction in the photosynthetic capacity of plants is presumed to negatively affect their fungal symbionts. To test this hypothesis under natural conditions, we artificially removed 100% of previous year needles in two successive years on Scots pine trees (Pinus sylvestris L.) to simulate pine sawfly attack. Despite a decline in the shoot growth of defoliated trees, root biomass did not differ from control trees. The ergosterol (fungal biomass) and starch concentration of fine roots, however, slightly declined in defoliated trees. Percent ectomycorrhizal colonization of fine root tips remained high in both defoliated and control trees. The dominant tubercle morphotypes were slightly more abundant in the control than in defoliated trees. In contrast to the relatively weak effects on vegetative ectomycorrhizae, reproduction declined near the defoliated pines. Average sporocarp numbers and, consequently, the relative fungal investment to reproduction of the estimated total fungal biomass were more than three times higher near controls than defoliated trees in the first treatment year. Defoliation also reduced the diversity of ectomycorrhizal species producing sporocarps. Mutualistic fungal symbionts may thus alter their reproductive investment in response to restrictions on host resources. Because fungal biomass in the roots as well as colonization percentage remained unchanged, Scots pine evidently continues to invest in the maintenance of the symbiosis despite the reduced photosynthetic capacity due to defoliation.

Defoliation increases carbon limitation in ectomycorrhizal symbiosis of Betula pubescens

Boreal forest trees are highly dependent on root-colonizing mycorrhizal fungi. Since the maintenance of mycorrhizal symbiosis implies a significant carbon cost for the host plant, the loss of photosynthetic leaf area due to herbivory is expected to reduce the host investment in mycorrhizae. We tested this hypothesis in a common garden experiment by exposing ectomycorrhizal white birch (Betula pubescens Ehrh.) seedlings to simulated insect defoliation of 50 or 100% intensity during either the previous or the current summer or repeatedly during both seasons before harvest. The shoot and root growth of the seedlings were distinctly reduced by both 100% defoliation and repeated 50% defoliation, and they were more strongly affected by previous-year than current-year defoliation. The root to shoot ratio significantly decreased after 100% defoliation, indicating reduced proportional allocation to the roots. Ergosterol concentration (i.e. fungal biomass) in the fine roots decreased by 100% defoliation conducted either in the year of harvest or in both years. No such decrease occurred following the 100% defoliation conducted in the previous year, indicating the importance of current photosynthates for fungal symbionts. The trend was similar in the colonization percentage of thick-mantled mycorrhizae in the roots, the most marked decline occurring in the repeatedly defoliated seedlings. The present results thus support the prediction that the plant investment in ectomycorrhizae may decline as a response to foliage loss. Moreover, the colonization percentage of thick-mantled mycorrhizae correlated positively with the ratio of leaf to heterotrophic plant biomass in the defoliated birch seedlings, but not in the control ones. This tends to indicate a stronger carbon limitation of ectomycorrhizal colonization in defoliated seedlings.

Defoliation-induced responses in peroxidases, phenolics, and polyamines in scots pine (Pinus sylvestris L.) needles

Effects of artificial defoliation on defensive needle chemistry in Scots pine (Pinus sylvestris L.) were evaluated with particular emphasis on peroxidases, phenolic compounds, soluble sugars, polyamines, and foliar nitrogen levels. The study was carried out on a nutrient-poor Scots pine stand with 8- to 25-year-old trees. Defoliation treatment consisted of repeated defoliation in two successive years with respective control trees. Defoliation was done before needle flushing by removing all mature needles. Guaiacol peroxidase activity increased in the needles after the first defoliation. The difference between treatments diminished towards autumn, and disappeared before the second defoliation in the next summer. After the second defoliation, the activities showed a similar trend. Apparently, peroxidases are involved in inducible chemical changes and recovery reactions that occur in the intact needles shortly after defoliation. After the second defoliation, total nitrogen concentration in the current year needles was about 20% lower, and free putrescine (a polyamine) concentration was 40% lower in the defoliated trees than in control needles. These changes indicate a loss of nitrogen due to defoliation. Specific phenolic compounds such as quercitrin, (+)-catechin, and two catechin derivatives increased in current year needles in response to defoliation. Accumulation of starch and sucrose in the current year needles of repeatedly defoliated trees may imply decreased assimilate transport. The results are indicative that changes in needle phytochemistry in response to defoliation accompany changes in needle nitrogen metabolism.

Induced accumulation of phenolics and sawfly performance in Scots pine in response to previous defoliation

Phenolic compounds often accumulate in foliar tissues of deciduous woody plants in response to previous insect defoliation, but similar responses have been observed infrequently in evergreen conifers. We studied the effects of defoliation on the foliar chemistry of Scots pine (Pinus sylvestris L.) and cocoon mass, and survival of the pine sawfly (Diprion pini L.). In two successive years, needles were excised early in the season leaving only the current-year shoot intact (defoliated trees); untreated entire shoots served as controls (control trees). A year after the second defoliation, pine sawfly larvae were transferred to the trees. Delayed induced resistance in Scots pine in response to defoliation was indicated by (1) reduced cocoon mass in defoliated trees and (2) increased concentrations of phenolics and soluble condensed tannins in the foliage of defoliated trees compared with controls. Myricetin-3-galactoside, which showed the strongest induced response (104% and 71% increase in current-year (C) and previous-year (C+1) needles) of the compounds analyzed, also entered the regression model explaining variation in sawfly performance. Other compounds that entered the model, e.g., (+)-catechin, showed weaker responses to defoliation than myricetin-3-galactoside. Hyperin, condensed tannins and quercitrin showed strong induced responses in C or C+1 needles, or both, but these compounds did not explain the variation in sawfly performance. Accumulation of phenolics is sometimes associated with the reduced foliage nitrogen (N) concentrations in deciduous trees, and our results suggest that this may also be the case in evergreen conifers. Based on the earlier findings that defoliation reduces needle N concentration and N deficiency results in the accumulation of the same phenolic compounds, i.e., myricetin and quercetin glycosides, and soluble condensed tannins, we suggest that the accumulation of phenolics in defoliated trees occurred in response to the reduced foliar N concentration.

Impact of wet deposited nickel on the cation content of a mat-forming lichen Cladina stellaris

The impact of experimentally sprayed aqueous nickel solution on the concentrations of potassium, calcium, magnesium and nickel in three horizontal strata (top, 0-20 mm; middle, 20-40 mm; and base, 40-60 mm) of the cushion-forming lichen Cladina stellaris was investigated. The experimental nickel deposition range used corresponded with that from the pristine forests of the Finnish border to polluted industrial sites of Russian Kola Peninsula (0-1000 mg Ni(2+) m(-2) year(-1)). The lichen mat retained ca. 31-66% of the nickel deposited during two growing seasons and the relative retention efficiency was highest at the low deposition end. The concentrations of cations in lichen thalli were significantly reduced only after the highest nickel deposition. Furthermore, the separate horizontal strata responded differently to nickel exposure indicating that the cation exchange sites of the top stratum were not completely saturated by nickel even after the most severe treatment. However, nickel deposited in high doses caused considerable reduction in potassium concentration indicating damage to cell membranes. Episodically deposited high concentrations of nickel can probably affect membrane integrity before detectable changes in total concentrations of cations in the lichen thallus take place. Thus, ratios of total concentrations of cations in the lichen thallus are fairly insensitive to nickel deposition, which reduces the risk of compounding effects when the ratios are used to indicate long-term acid deposition in areas with multiple pollution problems such as Kola Peninsula.

Does the lichen mat alleviate the effects of wet deposited nickel on soil microorganisms and Scots pine (Pinus sylvestris L.) seedlings

A field experiment was conducted in a dry heath forest dominated by Scots pine (Pinus sylvestris L.) and a mat-forming lichen (Cladina stellaris (Opiz) Brodo) to assess the effect of wet-deposited nickel (Ni) on pine seedlings and soil microorganisms, and to explore whether an intact lichen mat could act as a buffer against heavy metal deposits. Pine seedlings were planted in quadrats covered by a natural lichen layer and in quadrats from which the lichen layer had been completely removed. The quadrats were exposed to four levels of Ni deposition: 0 (i.e., distilled water), 10, 100 and 1000 mg m−2 year−1 in two growing seasons. Increasing Ni deposition led to an increase in the Ni content of the needles, roots and the soil organic layer. The lichen mat reduced Ni flow to the organic soil layer, but had no significant, reducing effect on needle or root Ni concentration. The most severe Ni treatment had detrimental effects on seedling growth and increased peroxidase activity in the previous years needles. Removal of the lichen layer did not increase susceptibility of seedlings to Ni. Values of maximal carbon use efficiency (Max) and metabolic quotient (qCO2) of the soil microorganisms indicated protective value of the lichen mat to soil microorganisms at the highest Ni treatment. Skimming per se decreased basal respiration, qCO2 and concentrations of potassium in the soil and also increased the lag period of the microorganisms as a response to in situ substrate addition.

Juuritutkimuksella uutta tietoa puiden kasvusta

Ilmastomme ominaispiirteitä ovat lämpötilan, valaistuksen ja sadannan voimakas vuodenaikaisvaihtelu. Ympäristötekijöiden vaihtelu edellyttää puiden maanpäällisiltä ja maanalaisilta kasvinosilta hyvää sopeutumiskykyä. Maanpäällisten osien kasvuun ja sopeutumiseen vaikuttavia tekijöitä on tutkittu laajasti. Sen sijaan puiden juuria on tutkittu vähemmän. Niiden merkitystä puiden kasvuun ei liene tunnistettu samassa määrin kuin maanpäällisten osien merkitystä. Juuristotutkimuksiin liittyy monia ongelmia. Näytteiden otto ja mittaukset ilman niiden kaivamista maasta ovat monin verroin hankalampia kuin mittaukset maanpäällisistä kasvinosista. Juurien toimintaa koskevaa tutkimustietoa kuitenkin tarvitaan, jotta voidaan ymmärtää puiden kasvuun vaikuttavia tekijöitä ja metsäekosysteemin ainevirtoja vaihtelevissa ympäristöoloissa. Viime vuosina tämän tutkimusalan tärkeys on ymmärretty paremmin ja monissa tutkimuslaitoksissa ja -laboratorioissa tutkimusta on kohdennettu myös puiden maanalaisten prosessien selvittämiseen. Muiden muassa Metsäntutkimuslaitoksella on Euroopan mittakaavassa ainutlaatuinen juuristolaboratorio http://www.metla. fi/jo/juuristolaboratorio/juuristolaboratorio.htm, joka perustettiin yhteistyössä Itä-Suomen yliopiston kanssa (kuva 1). Juurilla on merkittävä vaikutus metsäekosysteemien hiilenkiertoon. Arvioiden mukaan metsämaan osuus koko maaekosysteemin hiilivarastosta on 40 %, kun metsien maanpäällisten osien hiilivaraston osuus kaikesta maanpäällisten osien hiilestä on 80 %. Metsien hiilenkierron kannalta hienojuuret ovat lyhytikäisyytensä vuoksi merkittävä tekijä. Arvioiden mukaan jopa 75 % metsäekosysteemin biomassan nettotuotosta menee hienojuurten kasvuun. Eri puulajeja ja 28 maata käsittävässä kirjallisuuskatsauksessa todettiin, että sadanta lisää hienojuurten kasvua. Puun ikä lisää hienojuurten kasvua, mutta vähentää niiden uusiutumisnopeutta. Ympäristön stressitekijöiden vaikutuksista puiden juurten elintoimintoihin, kuolemaan, uusiutumisnopeuteen sekä juuristovaurioiden vaikutuksista maanpäällisten kasvinosien kasvuun tiedetään vähän.