Biomass distribution of two subalpine dwarf‐shrubs in relation to soil moisture and nutrient content

Question: Do soil water content and/or soil nitrogen (N) content and/or soil phosphorus (P) content affect the biomass of Vaccinium myrtillus and V. vitis‐idaea in a sub‐alpine heath? Location: Dolomites, northern Italy, 1800 m a.s.l. Methods: We determined above‐ground and below‐ground biomass of the shrubs at three sites, each on a different substrate type. At each site, we determined soil N‐ and P‐contents. We also determined leaf water potential (Psi;₁), N‐ and P‐concentrations in plant tissues and litter, as well as δ¹³C and δ¹⁵N in mature leaves. Results: V. myrtillus biomass was highest at the silicate site, V. vitis‐idaea biomass was highest at the carbonate site. Both shrubs had low biomass at the peat site, possibly due to a toxic effect of waterlogging in wet soils. For both species, pre‐dawn Psi;₁ indicated optimal hydration and midday Psi;₁ did not show any sign of water stress. Water use efficiency (WUE) did not differ among sites for any species. Whole‐plant nutrient concentrations showed that, with increasing biomass, N was diluted in V. myrtillus tissues while P was diluted in V. vitis‐idaea tissues. Foliar N‐concentration was higher overall for V. myrtillus. Foliar P‐concentration in V. myrtillus peaked at the silicate site. Foliar N : P ratios suggested that V. myrtillus was primarily P‐limited and V. vitis‐idaea primarily N‐limited. Conclusions: Water content affected the distribution of the two shrubs in a similar way, higher P‐availability in the soil enhanced V. myrtillus rather than V. vitis‐idaea.     

The shift in plant species composition in a subarctic mountain birch forest floor due to climate change would modify the biogenic volatile organic compound emission profile

Background and aims

Mountain birch forests dominate in the Subarctic but little is known of their non-methane biogenic volatile organic compound (BVOC) emissions. The dwarf shrubs Empetrum hermaphroditum, Vaccinium myrtillus and Vaccinium uliginosum co-dominate in the forest floors of these forests. The abundance of these three dwarf shrubs relative to each other could be affected by climate warming expected to increase nutrient availability by accelerating litter decomposition and nutrient mineralization. We 1) compared the BVOC emission profiles of vegetation covers dominated by E. hermaphroditum and V. myrtillus plus V. uliginosum in a subarctic mountain birch forest floor, 2) distinguished the BVOCs emitted from plants and soil and 3) measured how the BVOC emissions from the different vegetation covers differed under darkness.

Methods

BVOCs were sampled during two growing seasons using a conventional ecosystem chamber-based method, collected on adsorbent and analyzed with gas chromatography–mass spectrometry.

Results

High abundance of E. hermaphroditum increased the sesquiterpene emissions. Soil released fewer different BVOCs than controls (i.e. natural vegetation) but the total emission rates were similar. Darkness did not affect the emissions. Carbon emitted as BVOCs was less than 0.2% of the CO₂ exchange.

Conclusions

Our results suggest that sesquiterpene emissions from subarctic mountain birch forest floors would be reduced following an increased abundance of V. myrtillus and V. uliginosum with climate change because these species respond rapidly to increased nutrient availability.     

Tight Coupling Between Shoot Level Foliar N and P,Leaf Area,and Shoot Growth in Arctic Dwarf Shrubs Under Simulated Climate Change

Nutrient availability limits productivity of arctic ecosystems, and this constraint means that the amount of nitrogen (N) in plant canopies is an exceptionally strong predictor of vegetation productivity. However, climate change is predicted to increase nutrient availability leading to increases in carbon sequestration and shifts in community structure to more productive species. Despite tight coupling of productivity with canopy nutrients at the vegetation scale, it remains unknown how species/shoot level foliar nutrients couple to growth, or how climate change may influence foliar nutrients–productivity relationships to drive changes in ecosystem carbon gain and community structure. We investigated the influence of climate change on arctic plant growth relationships to shoot level foliar N and phosphorus (P) in three dominant subarctic dwarf shrubs using an 18-year warming and nutrient addition experiment. We found a tight coupling between total leaf N and P per shoot, leaf area and shoot extension. Furthermore, a steeper shoot length-leaf N relationship in deciduous species (Vaccinium myrtillus and Vaccinium uliginosum) under warming manipulations suggests a greater capacity for nitrogen to stimulate growth under warmer conditions in these species. This mechanism may help drive the considerable increases in deciduous shrub cover observed already in some arctic regions. Overall, our work provides the first evidence at the shoot level of tight coupling between foliar N and P, leaf area and growth i.e. consistent across species, and provides mechanistic insight into how interspecific differences in alleviation of nutrient limitation will alter community structure and primary productivity in a warmer Arctic.     

Understorey plant and soil responses to disturbance and increased nitrogen in boreal forests

Question: How do N fertilization and disturbance affect the understorey vegetation, microbial properties and soil nutrient concentration in boreal forests? Location: Kuusamo (66°22′N; 29°18′E) and Oulu (65°02′N; 25°47′E) in northern Finland. Methods: We conducted a fully factorial experiment with three factors: site (two levels), N fertilization (four levels) and disturbance (two levels). We measured treatment effects on understorey biomass, vegetation structure, and plant, soil and microbial N and C concentrations. Results: The understorey biomass was not affected by fertilization either in the control or in the disturbance treatment. Fertilization reduced the biomass of deciduous Vaccinium myrtillus. Disturbance had a negative effect on the biomass of V. myrtillus and evergreen Vaccinium vitis‐idaea and decreased the relative proportion of evergreen species. Fertilization and disturbance increased the biomass of grass Deschampsia flexuosa and the relative proportion of graminoids. The amount of NH₄⁺ increased in soil after fertilization, and microbial C decreased after disturbance. Conclusions: Our results suggest that the growth of slow‐growing Vaccinium species and soil microbes in boreal forests are not limited by N availability. However, significant changes in the proportion of dwarf shrubs to graminoids and a decrease in the biomass of V. myrtillus demonstrate the susceptibility of understorey vegetation to N enrichment. N enrichment and disturbance seem to have similar effects on understorey vegetation. Consequently, increasing N does not affect the rate or the direction of recovery after disturbance. Moreover, our study demonstrates the importance of understorey vegetation as a C source for soil microbes in boreal forests.     

The impacts of UV-B radiation on the regeneration of a sub-arctic heath community

The effects of ultraviolet-B radiation on regeneration after disturbance of a natural sub-arctic heathland have been investigated. Areas of pristine dwarf shrub heath were denuded of all above ground biomass in 1992 and exposed to enhanced UV-B (simulating a 15% depletion of the ozone layer). The resulting regenerated stem and leaf growth parameters were measured after four years on three dwarf shrubs, Vaccinium myrtillus, V. uliginosum and V. vitis-idaea and the grass Calamagrostis lapponica; leaves of the three dwarf shrubs were also analysed for UV-absorbing compounds and carbohydrates. Regeneration irrespective of treatment was slow, with Empetrum hermaphroditum failing to regenerate at all. Vaccinium myrtillus showed the most rapid regeneration attaining much of its original biomass in four years. There was a significant interaction between UV-B and year of regeneration in V. myrtillus; annual stem length increment showed an initial stimulation of 75% under enhanced UV-B in the first year of regeneration while a reduction of 16% was observed in the fourth year. Both V. uliginosum and V. vitis-idaea showed a reduction in annual stem length increment as regeneration progressed with a greater than 50% reduction in stem increment in the fourth year of regeneration compared to the first. Vaccinium uliginosum also showed an initial reduction in stem length increment of 40% under enhanced UV-B. None of the species were affected by enhanced UV-B in terms of total regenerated stem and leaf biomass or UV-absorbing compounds in regenerated leaf tissue. Total leaf carbohydrate and the ethanol/water soluble fraction in V. uliginosum were significantly increased by 29% and 31% respectively under enhanced UV-B. This suggests either a stimulation of photosynthesis or a reduction in sink size for photo-assimilates. Results are discussed in the context of the extremely slow regeneration of sub-arctic heath communities and the implications of contrasting UV-B effects on the regenerative ability of different species.     

Water- and nutrient-use efficiency of a deciduous species, Vaccinium myrtillus, and an evergreen species, V. vitis-idaea, in a subalpine dwarf shrub heath in the southern Alps, Italy

Periodic measurements of gas‐exchange rates and determinations of foliar N and P concentrations were used for evaluating instantaneous water‐use efficiency and photosynthetic nutrient‐use efficiency in two co‐existing dwarf shrubs of different growth form (V. myrtillus, deciduous, and V. vitis‐idaea, evergreen) in a subalpine heath in the southern Alps of Italy. Those data were compared with cumulative assessments of water‐use efficiency and photosynthetic nutrient‐use efficiency obtained by measuring leaf carbon isotope discrimination in leaf tissues and by estimating nutrient resorption from senescing leaves. V. myrtillus presented higher dry‐weight based rates of net photosynthesis (A_weight) compared to V. vitis‐idaea. A_weight was positively correlated with foliar‐nutrient status and intercellular‐to‐ambient gradient in CO₂ concentrations. A_weight was, furthermore, negatively correlated with leaf specific mass. Instantaneous photosynthetic nutrient‐use efficiency did not differ between the two species but the percentages of N and P pools resorbed from senescing leaves were somewhat higher in the deciduous species. The evergreen species showed lower P concentrations in senescing leaves which indicated a higher proficiency in resorbing phosphorus compared to the deciduous species. In addition, the evergreen species achieved a higher carbon gain per unit foliar N and P, due to a longer mean residence time of both nutrients. The two species did not differ from each other with respect to both instantaneous and long‐term water‐use efficiency. This was consistent with the climatic pattern, showing no sign of water deficiency through the growing season. Current‐year V. vitis‐idaea leaves had a significantly higher Δ¹³C compared to previous‐year leaves, possibly mirroring a long term acclimation of evergreen leaves, as far as they age, to the habitat conditions in the understory where evergreen species are usually confined within mixed dwarf‐shrub communities.     

长白山林线主要木本植物叶片养分的季节动态及回收效率

植物叶片养分含量的季节动态和回收效率对植被生态系统的养分循环和植物生长策略具有重要意义。以长白山高山林线上分布的3种主要木本植物——岳桦(Betula ermanii),牛皮杜鹃(Rhododendron aureum)和笃斯越橘(Vaccinium uliginosum)为研究对象,通过测定叶片中N、P、K、Ca、Mg、Fe等6种养分元素含量,分析在林线处植物叶片养分含量的季节动态及其与土壤养分含量的关系。结果表明岳桦和笃斯越橘叶片中养分元素的季节动态基本一致,即:N、P、K含量在生长季内逐渐降低,而Ca的含量逐渐增加;Mg在生长季旺盛期最低,而Fe含量却最高。牛皮杜鹃作为常绿灌木,叶片养分的季节动态与其余两种植物明显不同,表现为在生长季初期6种养分元素含量最低。岳桦和笃斯越橘植物叶片中N、P、K、Fe都有一定程度的回收,但笃斯越橘叶片的养分回收率更高,反映了笃斯越橘更能适应相对贫瘠的环境。3种林线植物叶片中养分含量与土壤养分并不存在显著的相关性,说明长白山林线上土壤中养分的分布没有对林线上3种主要的木本植物的生长和分布产生直接的影响。     

Growth and reproduction of the alpine grasshopper <Emphasis Type="Italic">Miramella alpina</Emphasis> feeding on CO<Subscript>2</Subscript>-enriched dwarf shrubs at treeline

The consequences for plant-insect interactions of atmospheric changes in alpine ecosystems are not well understood. Here, we tested the effects of elevated CO₂ on leaf quality in two dwarf shrub species (Vaccinium myrtillus and V. uliginosum) and the response of the alpine grasshopper (Miramella alpina) feeding on these plants in a field experiment at the alpine treeline (2,180 m a.s.l.) in Davos, Switzerland. Relative growth rates (RGR) of M. alpina nymphs were lower when they were feeding on V. myrtillus compared to V. uliginosum, and were affected by elevated CO₂ depending on plant species and nymph developmental stage. Changes in RGR correlated with CO₂-induced changes in leaf water, nitrogen, and starch concentrations. Elevated CO₂ resulted in reduced female adult weight irrespective of plant species, and prolonged development time on V. uliginosum only, but there were no significant differences in nymphal mortality. Newly molted adults of M. alpina produced lighter eggs and less secretion (serving as egg protection) under elevated CO₂. When grasshoppers had a choice among four different plant species grown either under ambient or elevated CO₂, V. myrtillus and V. uliginosum consumption increased under elevated CO₂ in females while it decreased in males compared to ambient CO₂-grown leaves. Our findings suggest that rising atmospheric CO₂ distinctly affects leaf chemistry in two important dwarf shrub species at the alpine treeline, leading to changes in feeding behavior, growth, and reproduction of the most important insect herbivore in this system. Changes in plant-grasshopper interactions might have significant long-term impacts on herbivore pressure, community dynamics and ecosystem stability in the alpine treeline ecotone.     

Impacts of multiple extreme winter warming events on sub‐Arctic heathland: phenology,reproduction, growth,and CO2 flux responses

Extreme weather events can have strong negative impacts on species survival and community structure when surpassing lethal thresholds. Extreme, short‐lived, winter warming events in the Arctic rapidly melt snow and expose ecosystems to unseasonably warm air (for instance, 2–10 °C for 2–14 days) but upon return to normal winter climate exposes the ecosystem to much colder temperatures due to the loss of insulating snow. Single events have been shown to reduce plant reproduction and increase shoot mortality, but impacts of multiple events are little understood as are the broader impacts on community structure, growth, carbon balance, and nutrient cycling. To address these issues, we simulated week‐long extreme winter warming events – using infrared heating lamps and soil warming cables – for 3 consecutive years in a sub‐Arctic heathland dominated by the dwarf shrubs Empetrum hermaphroditum, Vaccinium vitis‐idaea (both evergreen) and Vaccinium myrtillus (deciduous). During the growing seasons after the second and third winter event, spring bud burst was delayed by up to a week for E. hermaphroditum and V. myrtillus, and berry production reduced by 11–75% and 52–95% for E. hermaphroditum and V. myrtillus, respectively. Greater shoot mortality occurred in E. hermaphroditum (up to 52%), V. vitis‐idaea (51%), and V. myrtillus (80%). Root growth was reduced by more than 25% but soil nutrient availability remained unaffected. Gross primary productivity was reduced by more than 50% in the summer following the third simulation. Overall, the extent of damage was considerable, and critically plant responses were opposite in direction to the increased growth seen in long‐term summer warming simulations and the ‘greening’ seen for some arctic regions. Given the Arctic is warming more in winter than summer, and extreme events are predicted to become more frequent, this generates large uncertainty in our current understanding of arctic ecosystem responses to climate change.     

Bioassays of nutrient limitation in a tropical rain forest soil

Summary Six speices of shrubs and one large herb with contrasting life history patterns were used as bioassays of nutrient availability in a Costa Rican lowland rain forest soil. Growth responses of the herb (Phytolacca rivinoides, Phytolaccaceae) confirmed soil measurements indicating high availability of N and potentially limiting levels of P, K, Mg and Ca. Growth responses of the shrub species (Miconia spp., Melastomataceae and Piper spp., Piperaceae) to a complete nutrient fertilizer were generally less than that of Phytolacca. Lack of a strong shrub response to +P fertilization is probably due to mycorrhizal associations and slower growth rates of woody species. In general, increased growth did not occur at the expense of phenolic production in the leaves. The results emphasize that assessment of specific nutrient limitations to plant growth vary depending on species selected for the bioassay, even among species from the same community.     

Nitrogen-addition effects on leaf traits and photosynthetic carbon gain of boreal forest understory shrubs

Boreal coniferous forests are characterized by fairly open canopies where understory vegetation is an important component of ecosystem C and N cycling. We used an ecophysiological approach to study the effects of N additions on uptake and partitioning of C and N in two dominant understory shrubs: deciduous Vaccinium myrtillus in a Picea abies stand and evergreen Vaccinium vitis-idaea in a Pinus sylvestris stand in northern Sweden. N was added to these stands for 16 and 8 years, respectively, at rates of 0, 12.5, and 50 kg N ha^?1 year^?1. N addition at the highest rate increased foliar N and chlorophyll concentrations in both understory species. Canopy cover of P. abies also increased, decreasing light availability and leaf mass per area of V. myrtillus. Among leaves of either shrub, foliar N content did not explain variation in light-saturated CO₂ exchange rates. Instead photosynthetic capacity varied with stomatal conductance possibly reflecting plant hydraulic properties and within-site variation in water availability. Moreover, likely due to increased shading under P. abies and due to water limitations in the sandy soil under P. sylvestris, individuals of the two shrubs did not increase their biomass or shift their allocation between above- and belowground parts in response to N additions. Altogether, our results indicate that the understory shrubs in these systems show little response to N additions in terms of photosynthetic physiology or growth and that changes in their performance are mostly associated with responses of the tree canopy.     

Nitrogen response efficiency increased monotonically with decreasing soil resource availability: a case study from a semiarid grassland in northern China

The concept of nutrient use efficiency is central to understanding ecosystem functioning because it is the step in which plants can influence the return of nutrients to the soil pool and the quality of the litter. Theory suggests that nutrient efficiency increases unimodally with declining soil resources, but this has not been tested empirically for N and water in grassland ecosystems, where plant growth in these ecosystems is generally thought to be limited by soil N and moisture. In this paper, we tested the N uptake and the N use efficiency (NUE) of two Stipa species (S. grandis and S. krylovii) from 20 sites in the Inner Mongolia grassland by measuring the N content of net primary productivity (NPP). NUE is defined as the total net primary production per unit N absorbed. We further distinguished NUE from N response efficiency (NRE; production per unit N available). We found that NPP increased with soil N and water availability. Efficiency of whole-plant N use, uptake, and response increased monotonically with decreasing soil N and water, being higher on infertile (dry) habitats than on fertile (wet) habitats. We further considered NUE as the product of the N productivity (NP the rate of biomass increase per unit N in the plant) and the mean residence time (MRT; the ratio between the average N pool and the annual N uptake or loss). The NP and NUE of S. grandis growing usually in dry and N-poor habitats exceeded those of S. krylovii abundant in wet and N-rich habitats. NUE differed among sites, and was often affected by the evolutionary trade-off between NP and MRT, where plants and communities had adapted in a way to maximize either NP or MRT, but not both concurrently. Soil N availability and moisture influenced the community-level N uptake efficiency and ultimately the NRE, though the response to N was dependent on the plant community examined. These results show that soil N and water had exerted a great impact on the N efficiency in Stipa species. The intraspecific differences in N efficiency within both Stipa species along soil resource availability gradient may explain the differences in plant productivity on various soils, which will be conducive to our general understanding of the N cycling and vegetation dynamics in northern Chinese grasslands.     

Response patterns of Vaccinium myrtillus and V. vitis-idaea along nutrient gradients in boreal forest

Abstract. The relationships between biomass of dwarf shrub species and nutrient gradients of forest soils was studied under field conditions in boreal forests. The biomass-response curves of Vaccinium myrtillus and V. vitis-idaea were fitted against soil nutrient gradients using Generalized Linear Models (GLM). Ecological niches of Vaccinium myrtillus and V. vitis-idaea were evaluated, and effects of nitrogen addition (manipulation of the nutrient gradient) on response function were tested. The Vaccinium species showed statistically significant Gaussian responses along soil nitrogen, phosphorus and calcium gradients, but not along other gradients (K and Mg). Furthermore, manipulation of the nitrogen gradient seemed to have a minor effect on response functions, i.e. addition of nitrogen did not change ecological niches of these species. Ecological optima of V. myrtillus and V. vitis-idaea on the nutrient gradients were about the same. This study suggests that differences in dominance between Vaccinium myrtillus and V. vitis-idaea in boreal forest is not determined by nutrient gradients, but may rather be explained by light conditions and/or moisture availability.     

Inhibition of growth,and effects on nutrient uptake of arctic graminoids by leaf extracts — allelopathy or resource competition between plants and microbes?

Previous research has shown that plant extracts, e.g. from boreal dwarf shrubs and trees, can cause reduced growth of neighbouring plants: an effect known as allelopathy. To examine whether arctic and subarctic plants could also be affected by leaching of phytochemicals, we added extracts from the commonly occurring arctic dwarf shrubs Cassiope tetragona and Empetrum hermaphroditum, and from mountain birch, Betula pubescens ssp. tortuosa to three graminoid species, Carex bigelowii, Festuca vivipara and Luzula arcuata, grown in previously sterilized or non-sterilized arctic soils. The graminoids in non-sterilized soil grew more slowly than those in sterilized soil. Excised roots of the plants in non-sterilized soil had higher uptake rate of labelled P than those in sterilized soil, demonstrating larger nutrient deficiency. The difference in growth rate was probably caused by higher nutrient availability for plants in soils in which the microbial biomass was killed after soil sterilization. The dwarf shrub extracts contained low amounts of inorganic N and P and medium high amounts of carbohydrates. Betula extracts contained somewhat higher levels of N and much higher levels of P and carbohydrates. Addition of leaf extracts to the strongly nutrient limited graminoids in non-sterilized soil tended to reduce growth, whereas in the less nutrient limited sterilized soil it caused strong growth decline. Furthermore, the N and P uptake by excised roots of plants grown in both types of soil was high if extracts from the dwarf shrubs (with low P and N concentrations) had been added, whereas the P uptake declined but the N uptake increased after addition of the P-rich Betula extract. In contrast to the adverse extract effects on plants, soil microbial respiration and soil fungal biomass (ergosterol) was generally stimulated, most strongly after addition of the Betula extract. Although we cannot exclude the possibility that the reduced plant growth and the concomitant stimulation of microbial activity were caused by phytochemicals, we believe that this was more likely due to labile carbon in the extracts which stimulated microbial biomass and activity. As a result microbial uptake increased, thereby depleting the plant available pool of N and P, or, for the P-rich Betula extract, depleting soil inorganic N alone, to the extent of reducing plant growth. This chain of events is supported by the negative correlation between plant growth and sugar content in the three added extracts, and the positive correlation between microbial activity, fungal biomass production and sugar content, and are known reactions when labile carbon is added to nutrient deficient soils.     

Responses of ground vegetation species to clear-cutting in a boreal forest: aboveground biomass and nutrient contents during the first 7 years

Rapid growth of ground vegetation following clear-cutting is important to site productivity because vegetation retains nutrients in the ecosystem and can decrease nutrient leaching prior to stand re-establishment. Aboveground biomass, nutrient contents (N, P, K and Ca) and species composition of ground vegetation were determined 1 year before and for 7 years after clear-cutting of a mixed forest dominated by Norway spruce [Picea abies (L.) H. Karst.] in eastern Finland. The biomass of the feather mosses [Pleurozium schreberi Brid. and Hylocomium splendens (Hedw.) B. S.& G.] and the dwarf shrubs (Vaccinium myrtillus L. and V. vitis-idaea L.), which had dominated the ground vegetation in the mature forest, significantly decreased after clear-cutting. However, with the exception of H. splendens, these species had recovered within 3–5 years. The biomass of Deschampsia flexuosa (L.) Trin. considerably increased soon after clear-cutting, and Epilobium angustifolium L. appeared 3–5 years after cutting. These species contributed to the retention of nutrients not simply because of their biomass but also because of higher nutrient concentrations in their tissues. Total biomass and nutrient contents of the ground vegetation exceeded those of the pre-cutting levels. The proportion of ground vegetation biomass and nutrient contents represented by mosses decreased after cutting, while V. myrtillus, although reduced after cutting, remained a marked nutrient sink. The results suggest that H. splendens is the most sensitive species to cutting, but the biomass of P. schreberi, V. myrtillus and V. vitis-idaea return to initial levels soon after clear-cutting as do the nutrient contents of ground vegetation.     

Nutrient limitation in boreal rich‐fen vegetation: A fertilization experiment

Abstract. Rich‐fen vegetation influenced by hay‐making in the Sølendet Nature Reserve, Central Norway, was fertilized with N, P and K in a full‐factorial fertilization experiment to investigate the nutrient limitation of plant growth at both community and species levels. Above‐ground biomass, shoot density and nutrient concentration were measured in several species and groups of species at three sites after two years of fertilization. At the community level, the results indicate multiple limitation by N and P in the two least productive rich‐fen communities: one characterized by small sedges and herbs, and the other by high abundance of Menyanthes trifoliata and tall sedges. Increased nutrient availability had no effect on a more highly productive, tall‐growing, spring‐influenced community, indicating no nutrient limitation. The results at the species level correspond well with those at the community level, indicating multiple limitation by N and P in most of the dominant and sub‐dominant species. However, P seems to limit growth more than N in Succisa pratensis, and N seems to limit growth more than P in Carex panicea. Furthermore, Eriophorum angustifolium seems to be limited by K. The results did not show which nutrient limits the growth of Carex dioica, C. lasiocarpa and Trichophorum cespitosum. Indications that growth in low‐productive, boreal rich‐fen communities is generally limited by P was not confirmed.     

Growth and allocation of the arctic sedges Eriohorum angustifolium and E. vaginatum: effects of variable soil oxygen and nutrient availability

In arctic tundra soil, oxygen depletion associated with soil flooding may control plant growth either directly through anoxia or indirectly through effects on nutrient availability. This study was designed to evaluate whether plant growth and physiology of two arctic sedge species are more strongly controlled by the direct or indirect effects of decreased soil aeration. Eriophorum angustifolium and E. vaginatum, which originate from flooded and well-drained habitats, respectively, were grown in an in situ transplant garden at two levels of soil oxygen, nitrogen, and phosphorus availability over two growing seasons. In both species, N addition had a stronger effect on growth and biomass allocation than P addition or soil oxygen depletion. Net photosynthesis and carbohydrate concentrations were relatively insensitive to changes in these factors. Biomass reallocated from shoots to below-ground parts in response to limited N supply was equally divided between roots (nutrient acquisition) and perennating rhizomes (storage tissue formation) in E. angustifolium. E. Vaginatum only increased its allocation to rhizomes. In the flood-tolerant E. angustifolium, growth was improved by soil anoxia and biomass allocation among plant parts was not significantly affected. Contrary to our initial hypothesis, whole-plant growth in E. vaginatum improved in flooded soils; however, it only did so when N availability was high. Under low N availability growth in flooded soils was reduced by 20% compared to growth in the aerobic environment. Reduced biomass allocation to rhizomes and thus to storage potential under anaerobic conditions may reduce long-term survival of E. vaginatum in flooded habitats.     

Nutrient relocation,hydrological functions,and soil chemistry in plantations as compared to natural forests in central Yunnan,China

The relocation of nutrients among dominant plant species, along with hydrological functions and soil chemistry in five plant communities, including Eucalyptus plantation, Pinus plantation, shrubland, semi-natural, and natural secondary forests were investigated in central Yunnan, China. The nutrient P, N, and K accumulation in above-ground biomass of Eucalyptus smithii (stems, barks, branches, and leaves) were the highest, followed by Pinus yunnanensis of both the Pinus plantation and the semi-natural forest. The nutrient retranslocation efficiency (NRE) of E. smithii was the highest for nutrient P, N, and K with values of 56, 66, and 67%, respectively, among the dominant plant species of the five plant communities, while the NRE of P. yunnanensis in Pinus plantation had the second highest value of NRE for nutrient N. The nutrient content (available P, N, and soil organic matter) in the upper two soil layers under Eucalyptus and Pinus plantations was correspondingly found to be lower than that of the other forests. Moreover, under the Eucalyptus and Pinus plantations, surface runoff, soil erosion, and nutrient loss were more serious, and the water storage of litterfall and canopy interception were significantly lower than that in the other plant communities. Accordingly, we suggest that single-species plantations cannot present the same ecological benefits as natural forests, because of their simple, uniform structures, and the characteristics of the dominant plant species.     

Pollen limitation in three sympatric species ofVaccinium (Ericaceae) in the Upper Ardennes, Belgium

The reproductive success of three co-flowering species ofVaccinium (V. myrtillus, V. vitis-idaea andV. uliginosum) was studied in one heathland of the Upper Ardennes, Belgium, during three years (1988–1990). The purpose was to examine whether pollen limitation, flower position and flowering phenology may influence patterns of fruit set in these three sympatric species. I quantified fruit and seed set following supplementary hand-pollinations and compared this to natural fruit set. On the same plants, I also quantified fruit and seed set in relation to the spatial position of the flowers on the ramet and their temporal sequence of blooming. Hand-pollination had no significant effect on fruit set inV. vitis-idaea andV. uliginosum, but significantly increased seed number per fruit in 1989. InV. myrtillus both fruit and seed set were increased by supplementary pollination, but significantly in only one year. Analyses of position effects revealed that the fruits in the lower positions in the ramet did not mature preferentially and did not contain more seeds inV. uliginosum and inV. vitis-idaea. Flowering phenology also had no significant effect for these species. InV. myrtillus hand-pollinated flowers showed a seasonal decline in seed number, and control (naturally pollinated) flowers showed a seasonal increase in fruit set. Fruit and seed set appear to be pollen-limited rather than resource-limited inV. uliginosum. In the two other species, unfavorable weather (frosts) could be a more important cause of low fruit and seed set.     

Seasonal grazing effects by semi-domesticated reindeer on subarctic mountain birch forests

In northern Fennoscandia, the spatial and temporal grazing practices of semi-domesticated reindeer (Rangifer tarandus tarandus) vary, which implies different grazing effects dependent on natural conditions as well as management regime (i.e., timing and intensity of grazing). We compared density and biomass of main plant groups in semi-dry mountain birch forests exposed to either long-term summer or winter grazing in three reindeer herding districts in the northernmost Finland. Percent plant cover, height, and biomass of reindeer lichens (Cladonia spp.) and dwarf shrubs (Vaccinium uliginosum, Calluna vulgaris, and Betula nana) were lower on summer ranges compared with winter ranges. The biomass of other dwarf shrubs (Vaccinium myrtillus and V. vitis-idaea), and graminoids and herbs, and the % cover of non-vegetated bare soil and litter were, however, higher on summer ranges than on winter ranges. Young mountain birch shoots (Betula pubescens ssp. czerepanovii) were less frequent on summer ranges than on winter ranges. The total leaf biomass under the browsing height of reindeer (<1.5 m) was also lower on summer ranges compared with winter ranges. Especially in drier and nutrient poor mountain birch forests, intensive summer grazing reduces the quantity of lichens and total plant biomass which affects the ecological state and productivity of these forests and also reduces especially their winter grazing value for reindeer. Therefore, in addition to regulating the maximum sustained numbers of reindeer, pasture rotation systems that effectively protect dry and nutrient poor vegetation from summer grazing and trampling should be encouraged.