Françoise Martz

Phenolic Composition and Antioxidant Capacity of Bilberry (Vaccinium myrtillus) Leaves in Northern Europe Following Foliar Development and Along Environmental Gradients

Bilberry is a characteristic field layer species in the boreal forests and is an important forage plant for herbivores of the North European ecosystem. Bilberry leaves contain high levels of phenolic compounds, especially hydroxycinnamic acids, flavonols, catechins, and proanthocyanidins. We investigated the phenolic composition of bilberry leaves in two studies, one following foliar development in forest and open areas, and the other along a wide geographical gradient from south to north boreal forests in Finland. An analysis of bilberry leaves collected in open and forest areas showed that major phenolic changes appeared in the first stages of leaf development, but, most importantly, synthesis and accumulation of flavonoids was delayed in the forest compared to the high light sites. Sampling along a geographical gradient in the boreal zone indicated that leaves from higher latitudes and higher altitudes had greater soluble phenolic and flavonol levels, higher antioxidant capacity, and lower contents of chlorogenic acid derivatives. The ecological significance of the results is discussed.

Does climate change influence the availability and quality of reindeer forage plants

The northward and upward movement of the tree line and gradual replacement of lichens with vascular plants associated with increasing temperatures and nutrient availability may change the reindeer pastures in Northern Fennoscandia. The productivity of reindeer forage will most probably increase, but their protein (nitrogen) concentrations may decrease because of higher temperatures and CO2 concentration. In the long term, the nutritive value of forage will depend on the mineralization rate and nutrient uptake from the soil. Enhanced UV-B is likely to increase the concentration of phenolics, decreasing forage quality and choice, but reindeer may adapt to increased phenolics. Increased winter precipitation, the occurrence of ice layers, deeper snow cover, and the appearance of molds beneath the snow cover may reduce the availability and/or quality of reindeer forage, but prolongation of snowless periods might have the opposite effect. The net balance of negative and positive effects will vary regionally depending on the climate, bedrock, vegetation, reindeer herding systems and socio-political factors. Multidisciplinary research is needed most importantly on the effects of the changing winter climate on reindeer forage, and the effect of modified forage quality on reindeer physiology.

Phenolic Responses of Mountain Crowberry (Empetrum nigrum ssp. hermaphroditum) to Global Climate Change are Compound Specific and Depend on Grazing by Reindeer (Rangifer tarandus)

Mountain crowberry (Empetrum nigrum ssp. hermaphroditum) is a keystone species in northern ecosystems and exerts important ecosystem-level effects through high concentrations of phenolic metabolites. It has not been investigated how crowberry phenolics will respond to global climate change. In the tundra, grazing by reindeer (Rangifer tarandus) affects vegetation and soil nutrient availability, but almost nothing is known about the interactions between grazing and global climate change on plant phenolics. We performed a factorial warming and fertilization experiment in a tundra ecosystem under light grazing and heavy grazing and analyzed individual foliar phenolics and crowberry abundance. Crowberry was more abundant under light grazing than heavy grazing. Although phenolic concentrations did not differ between grazing intensities, responses of crowberry abundance and phenolic concentrations to warming varied significantly depending on grazing intensity. Under light grazing, warming increased crowberry abundance and the concentration of stilbenes, but decreased e.g., the concentrations of flavonols, condensed tannins, and batatasin-III, resulting in no change in total phenolics. Under heavy grazing, warming did not affect crowberry abundance, and induced a weak but consistent decrease among the different phenolic compound groups, resulting in a net decrease in total phenolics. Our results show that the different phenolic compound groups may show varying or even opposing responses to warming in the tundra at different levels of grazing intensity. Even when plant phenolic concentrations do not directly respond to grazing, grazers may have a key control over plant responses to changes in the abiotic environment, reflecting multiple adaptive purposes of plant phenolics and complex interactions between the biotic and the abiotic factors.

Effect of the temperature and the exclusion of UVB radiation on the phenolics and iridoids in Menyanthes trifoliata L. leaves in the subarctic

The long-term effects of UVB exclusion and temperature on the methanol extractable (ME) phenolics (flavonoids, phenolic acids) and iridoids of Menyanthes trifoliata L. (Mt) leaves were studied in northern Finland (68 degrees N) using wooden frames covered with filters for UVB exclusion (polyester filter), control (cellulose acetate filter) and ambient (no filter) conditions. Analysis of ambient plots showed no effect of the daily mean temperature (2sigma = 1.58 degrees C) on the leaf ME compound content and composition, but minimum temperatures decreased the flavonol content. UVB exclusion did not affect the total ME compound content but significantly decreased the proportion of flavonols concomitantly with an increase in iridoids. Due to its high iridoid content, Mt appears as an interesting model plant for studying the iridoid biosynthesis and its regulation under stress conditions.

The Snow Must Go On: Ground Ice Encasement, Snow Compaction and Absence of Snow Differently Cause Soil Hypoxia, CO2 Accumulation and Tree Seedling Damage in Boreal Forest.

At high latitudes, the climate has warmed at twice the rate of the global average with most changes observed in autumn, winter and spring. Increasing winter temperatures and wide temperature fluctuations are leading to more frequent rain-on-snow events and freeze-thaw cycles causing snow compaction and formation of ice layers in the snowpack, thus creating ice encasement (IE). By decreasing the snowpack insulation capacity and restricting soil-atmosphere gas exchange, modification of the snow properties may lead to colder soil but also to hypoxia and accumulation of trace gases in the subnivean environment. To test the effects of these overwintering conditions changes on plant winter survival and growth, we established a snow manipulation experiment in a coniferous forest in Northern Finland with Norway spruce and Scots pine seedlings. In addition to ambient conditions and prevention of IE, we applied three snow manipulation levels: IE created by artificial rain-on-snow events, snow compaction and complete snow removal. Snow removal led to deeper soil frost during winter, but no clear effect of IE or snow compaction done in early winter was observed on soil temperature. Hypoxia and accumulation of CO2 were highest in the IE plots but, more importantly, the duration of CO2 concentration above 5% was 17 days in IE plots compared to 0 days in ambient plots. IE was the most damaging winter condition for both species, decreasing the proportion of healthy seedlings by 47% for spruce and 76% for pine compared to ambient conditions. Seedlings in all three treatments tended to grow less than seedlings in ambient conditions but only IE had a significant effect on spruce growth. Our results demonstrate a negative impact of winter climate change on boreal forest regeneration and productivity. Changing snow conditions may thus partially mitigate the positive effect of increasing growing season temperatures on boreal forest productivity.

Different response of two reindeer forage plants to enhanced UV-B radiation: modification of the phenolic composition

The long-term effects of enhanced UV-B radiation on the content and composition of leaf phenolics in Epilobiumangustifolium L. and Eriophorumrusseolum Fries ex Hartman were studied in northern Finland (68°N) using two UV-B enhancement experiments, both simulating UV-BCIE radiation and corresponding to a 20% loss of ozone layer. High proportions of hydrolyzable tannins (69%) and condensed tannins (66%) characterized both Epilobium and Eriophorum leaves, respectively. No UV treatment effect was detected in the content or composition of Epilobium leaf soluble phenolics, whereas significant UV effects were detected in Eriophorum leaves in a developmental-specific manner. At the end of the growing season, the proportion of total soluble phenolics was higher in leaves exposed to enhanced UV-A and UV-B radiation than in the control leaves, but the phenolic composition was not significantly modified. This study introduces a new example on plants’ phenolic response to UV radiation being species-specific and detectable only at certain developmental stages. Possible consequences of increased phenolic content in forage plants for selection and digestibility by reindeer are, however, not yet known.

Bacterial and fungal communities in boreal forest soil are insensitive to changes in snow cover conditions

The northern regions are experiencing considerable changes in winter climate leading to more frequent warm periods, rain-on-snow events and reduced snow pack diminishing the insulation properties of snow cover and increasing soil frost and freeze-thaw cycles. In this study, we investigated how the lack of snow cover, formation of ice encasement and snow compaction affect the size, structure and activities of soil bacterial and fungal communities. Contrary to our hypotheses, snow manipulation treatments over one winter had limited influence on microbial community structure, bacterial or fungal copy numbers or enzyme activities. However, microbial community structure and activities shifted seasonally among soils sampled before snow melt, in early and late growing season and seemed driven by substrate availability. Bacterial and fungal communities were dominated by stress-resistant taxa such as the orders Acidobacteriales, Chaetothyriales and Helotiales that are likely adapted to adverse winter conditions. This study indicated that microbial communities in acidic northern boreal forest soil may be insensitive to direct effects of changing snow cover. However, in long term, the detrimental effects of increased ice and frost to plant roots may alter plant derived carbon and nutrient pools to the soil likely leading to stronger microbial responses.