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.     

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.     

Experimental studies on Vaccinium myrtillus and Vaccinium vitis-idaea in relation to air pollution and global change at northern high latitudes: A review

The aim of this paper is to review the experimental studies performed with two important species (i.e. Vaccinium myrtillus and Vaccinium vitis-idaea) of the shrub layer of boreal forests. The focus is on ecophysiology and stress tolerance under conditions of air pollution and global change. Our objective was to make an overview of studies of abiotic stress related to global change on the above species, and discuss the reported effects of these environmental factors. These issues include nitrogen, heavy metals, radionuclides, salt, ozone, carbon dioxide, warming climate, declining snow cover, periodic droughts, fire and elevated UV radiation. The findings suggest that both species are relatively tolerant to many abiotic environmental stresses: increased nitrogen and metal supply have negligible impacts on these species, as the atmospheric gaseous compounds reported. In some cases the effects are even positive, since extra nitrogen may improve the frost hardiness of V. vitis-idaea, and V. myrtillus may even benefit from elevated CO₂. Both species also seem to recover well from fires. However, the reports demonstrated that both have species-specific areas of weakness: (1) V. myrtillus is susceptible to stress caused by warming winter, and (2) frost hardiness of V. vitis-idaea may be reduced under enhanced UV radiation.     

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.     

Niche differentiation with respect to light utilization among coexisting dwarf shrubs in a subarctic woodland

Summary Canopy structure, shoot design, and photosynthetic light recruitment were used to compare four coexisting dwarf shrub species with respect to light utilization. All four species showed different shoot designs which probably result in different light interception properties. Leaves of Vaccinium uliginosum showed the highest levels of photosynthetic light saturation but in situ the shoots of this species reached their maximum photosynthetic rate at the lowest photon flux densities. No consistent differences with respect to photosynthetic light responses were found between deciduous and evergreen species. At sites dominated by one of the deciduous species (Vaccinium uliginosum or V. myrtillus), the two evergreen species studied (V. vitis-idaea and Empetrum hermaphroditum) occurred in the understory, i.e., with their leaf distribution slightly below that of the deciduous species. Sites dominated by one of the evergreen species showed less vertical differentiation in leaf distribution between species.     

Effects of leaf nitrogen availability and leaf position on nitrogen allocation patterns in Vaccinium vitis-idaea and Vaccinium uliginosum

Summary Nitrogen allocation patterns from leaves of Vaccinium vitis-idaea (evergreen) and Vaccinium uliginosum (deciduous) were assessed using a foliar application of ¹⁵N labeled ammonium sulfate. These are wild perennial shrubs inhabiting arctic and subarctic regions. More label was transported from labeled leaves of Vaccinium uliginosum then Vaccinium vitis-idaea. In Vaccinium uliginosum, the amount of label transported from the labeled leaf increased as the concentration of nitrogen in the label increased. Current growth in Vaccinium uliginosum was a strong sink for nitrogen because most of the ¹⁵N transported from the labeled leaf was contained in this region. In addition, when greater quantities of nitrogen were applied, larger quantities were retained in current growth. Current growth of Vaccinium vitis-idaea, on the other hand, was not as strong a sink because regardless of the nitrogen available thru various label concentrations, the enrichment of current growth was not affected and was not significantly different from older stems or leaves. Yet, in both species, nitrogen was transported freely from leaves of all positions along the stem to all parts of the plant including roots and rhizomes. The position of the leaf along the stem had no effect on the patterns of allocation to other organs.     

Cytological observations on some species and hybrids ofVaccinium

Summary Meiosis was studied inVaccinium myrtillus (n=12),V. vitis-idaea (n=12),V. myrtillus ×vitisidaea (n=12), colchicine-induced tetraploidV. myrtillus (n=24),V. uliginosum (n=24), the cultivated blueberry varieties Rancocas and Pemberton (n=24), and the hybridsV. uliginosum × Rancocas andV. uliginosum × Pemberton (n=24). Pairing was regular in the diploid species. Some multivalent formations occurred inV. uliginosum and in the cultivated blueberry varieties and to a considerable extent in the autotetraploidV. myrtillus. The spontaneous diploid hybridV. myrtillus ×vitis-idaea displayed numerous meiotic irregularities, whereas the artificial hybrids betweenV. uliginosum and cultivated blueberry showed relatively regular meiosis. Pairing relationships of the various genomes are discussed. Breeding programs for the use of North-European species with American cultivated blueberry varieties are discussed in the light of the cytological observations.

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Zusammenfassung Die diploiden (n=12) ArtenVaccinium myrtillus, V. vitis-idaea, V. myrtillus×vitis-idaea und die tetraploiden (n=24)V. myrtillus (Colchicin-induziert) undV. uliginosum sowie die kultivierten amerikanischen Blaubeersorten Rancocas und Pemberton (n=24) und die HybridenV. uliginosum×Rancocas undV. uliginosum×Pemberton wurden cytologisch untersucht. In den diploiden Arten war die meiotische Chromosomenpaarung regelmäßig. Multivalente traten in geringerem Umfange beiV. uliginosum und kultivierten Blaubeersorten, viel stärker aber in der autotetraploidenV. myrtillus auf. Die spontane diploide HybrideV. myrtillus×vitis-idaea zeigte zahlreiche meiotische Störungen, die künstlichen Hybriden zwischenV. uliginosum und der kultivierten amerikanischen Blaubeere hatten demgegenüber eine relativ regelmäßige Meiose. Die Paarungsverhältnisse der verschiedenen Genome werden diskutiert. Auf Grund der cytologischen Beobachtungen wird ein Züchtungsprogramm für nordeuropäischeVaccinium-Arten zusammen mit amerikanischen Kultursorten erörtert.

     

Phenolic compounds in berries and flowers of a natural hybrid between bilberry and lingonberry (Vaccinium × intermedium Ruthe)

Hybridization between species plays an important role in the evolution of secondary metabolites and in the formation of combinations of existing secondary metabolites in plants. We have investigated the content of phenolic compounds in berries and flowers of Vaccinium × intermedium Ruthe, which is a rare natural hybrid between bilberry (Vaccinium myrtillus L.) and lingonberry (Vaccinium vitis-idaea L.). The berries and flowers of the hybrid showed characteristics inherited from both parent species in the distribution and contents of phenolic compounds. Bilberry is known as one of the richest sources of anthocyanins and to have a profile of 15 major forms combining cyanidin, delphinidin, petunidin, peonidin and malvidin with galactose, glucose and arabinose. Lingonberry contains only cyanidin glycosides. Hybrid berries contained all bilberry anthocyanins with pronounced cyanidin content. With regard to proanthocyanidins and flavonol glycosides, the hybrid inherited diverse profiles combining those of both parental species. The distribution of hydroxycinnamic acids was quite uniform in all studied berries. Of the identified compounds, 30 were detected in lingonberry, 46 in bilberry, 53 in hybrid berries and 38 in hybrid flowers. Hence, compared with the parent species, hybrid berries possess a more diverse profile of phenolic compounds and, therefore, can offer interesting material for breeding purposes.     

Growth performance of two deciduous Vaccinium species in relation to nutrient status in a subalpine heath

Net primary production (NPP) of two Vaccinium species (V. myrtillus and V. uliginosum) was determined in three subalpine heath communities on the Northern Apennines (N. Italy). The main objective of the study was to test the hypothesis that two species sharing the same plant functional type (deciduous dwarf shrub) have similar growth performances along environmental gradients. The second objective was to assess whether, and to what extent, NPP of the two species was associated with functional and morphological traits, which can affect plant growth in relation to nutrient status. Total community NPP in the three communities was closely related to soil nutrient availability. NPP of V. uliginosum did not vary among communities, while that of V. myrtillus peaked in the most fertile habitat. The N:P ratio in the whole plant as well as in the leaves of the two shrubs exactly mirrored the among-community pattern in soil phosphate concentration. In particular, the foliar N:P ratio in both V. uliginosum and V. myrtillus was >16 in the poorer sites, which indicates P limitation. I concluded that the growth response of the two shrubs in relation to soil nutrient availability is individualistic. Growth of V. myrtillus is P-limited while that of V. uliginosum is not.     

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.     

Effect of industrial air pollution on wild plant seed germination and seedling growth

Germination of the following wild plant seeds was studied: bilberry (Vaccinium myrtillus L.), cowberry (V. vitis-idaea L.), bog bilberry (V. uliginosum L.), black crowberry (Empetrum hermaphroditum Hagerup), bearberry (Arctostaphylos uva-ursi (L.) Spreng.), bunchberry (Chamaepericlymenum suecicum (L.) Aschers. et Graebn.), cottongrass (Eriophorum polystachion L.), goldenrod (Solidago lapponica With.), fireweed (Chamaenerion angustifolium (L.) Scop.), marsh cinquefoil (Comarum palustre L.), cloudberry (Rubus chamaemorus L.), and Scots pine (Pinus sylvestris L.). The seeds were collected at different distances from the source of industrial air pollution (Severonickel smelter complex, Murmansk region). The task was the assessment of potential possibility of restoring corrupted north forest plant communities via seed propagation. By the response of reproductive structures to stressor, investigated species were divided into 3 groups: tolerant (members of the genus Vaccinium and Ch. angustifolium); moderately tolerant (C. palustre, E. polystachion, A. uva-ursi, and R. chamaemorus); and sensitive (P. sylvestris, E. hermaphroditum, and S. lapponica). Laboratory seed germinability of Vaccinium species was high (> 90%) regardless of the levels of Ni and Cu accumulation in the seeds and the index of technogenic load, whereas this index in E. hermaphroditum and P. sylvestris seeds was significantly reduced with the increase in the heavy metal contents in the seeds. The greenhouse experiments with the seeds of three Vaccinium species collected in the background area and in the zone of the highest pollution and germinated on the forest litter from the same sites and observation for seedling development allow us to conclude that a potential possibility of these species to seed propagation are not limited by their seed viability even under conditions of the highest technogenic load. In sites of environment pollution, the high metal content in the upper soil layer is the main factor limiting plant seed propagation.     

Lycopodium annotinum and light quality: Growth responses under canopies of twoVaccinium species

Lycopodium annotinum, a clonal pteridophyte, was grown under canopies formed by various combinations of deciduousVaccinium myrtillus and evergreenV. vitis-idaea. The canopies differed in red:far-red ratio, withV. myrtillus giving the lowest ratio. The current year's horizontal segments ofL. annotinum had the same specific weight in all treatments, but the lengths of the segments were significantly increased when grown under the canopy ofV. myrtillus compared withV. vitis-idaea. The number of vertical apices was also increased. Both these responses increased the amount of assimilating structures ofL. annotinum when growing under aV. myrtillus canopy. This might be beneficial for the evergreenL. annotinum, since it can then utilize spring and autumn periods when the deciduousV. myrtillus has no leaves.     

Photosynthetic characteristics and leaf carbon economy of a deciduous and an evergreen dwarf shrub: Vaccinium uliginosum L. and V. vitis-idaea L.

The seasonal course of photosynthetic rate, and light and temperature relations were studied in the dwarf shrubs Vaccinium uliginosum L., deciduous, and Vaccinium vitisidaea L., evergreen, at a subarctic site in northern Sweden, Using the photosynthetic characteristics and meteorological data from the site, the seasonal and life-span carbon dioxide gain was estimated. The photosynthetic capacity of V. uliginosum was at a maximum one month after the start of leaf expansion and declined rapidly in the beginning of September. The old V. vitis-idaea leaves needed about 2 wk to recover full photosynthetic capacity after snow-melt; the current-year V. vitis-idaea leaves needed the same time after bud-break to reach full capacity. The leaves of V. vitis-idaea showed no seasonal trend in photosynthetic capacity after the first two wk of recovery, but their capacity decreased by one third after the first winter and by approximately 10% yr^?1 over the following two yr. The seasonal variation in the photosynthetic response to temperature was more marked in V. uliginosum than in V. vitis-idaea. Light saturation occurred at approximately 3000 μmol m^?2 s^?1 in V. uliginosum and at 60 μmol m^?2 s^?1 in one-year-old V. vitis-idaea leaves. The leaves of both species had a positive carbon balance at photon flux densities above 5 μmol m^?2 s^?1. The calculated seasonal CO² gain was 21 g CO₂ g_?1 leaf in V. uliginosum and 6–8 g CO₂ g_?1 in V. vitis-idaea leaves. Life-span CO₂ gain for leaves of V. vitis-idaea was the same as in V. uliginosum, viz. 21 g CO₂ g_?1. One fifth of the CO₂ gain of V. vitis-idaea was assimilated during periods when V. uliginosum was leafless.     

植物冷驯化相关信号机制

植物经过非致死温度的处理可以获得更强的抗冷能力叫做冷驯化,主要包括寒驯化和冻驯化 .在冷驯化过程中,质膜首先感受冷信号,调节胞质中IP3的含量,诱导胞质Ca2+浓度的升高,从而激活CBF基因的表达.至今已经克隆了大量的冷调控基因,组成了复杂的信号传导网络,其中ICE1-CBF-COR通路在植物的冷驯化过程中起到重要的作用.ICE1基因编码一个MYB类型的碱性螺旋 环-螺旋（bHLH）转录因子,在上游调节CBF和 其它转录因子的表达,提高抗冷性. HOS1蛋白通过泛素化介导的蛋白降解负调控ICE1,另外,CBF还通过转录的自我调控保持恰当的表达水平.基因的分析研究证明,RNA修饰和核质转运在植物的抗冷过程中也具有重要作用.在不依赖于CBF的途径中,转录因子HOS9和HOS10在调节抗冷有关基因的表达和提高抗冷能力方面具有至关重要的作用.     

The functional quality of decomposing litter outputs from an Arctic plant community is affected by long-term exposure to enhanced UV-B

Plant exposure to enhanced UV-B radiation typically induces changes in leaf secondary metabolite profiles which will be inherited in litter, affecting litter breakdown and the carbon (C) dynamics of sensitive plant communities. A key enzyme in the decomposition process is phenol oxidase which is influenced by litter quality and, hence, a decomposition bioindicator. Here we investigated dwarf shrub litter decomposition following experimental community exposure to enhanced UV-B over two decades in the Swedish sub-Arctic. We examined the hypothesis that foliar UV-B exposure would alter litter quality to elevate phenol oxidase activity. This was tested in the field by measuring phenol oxidase activity in freshly collected mixed-community litter from under our experimental vegetation. A laboratory mesocosm was next used in a decomposition assay to investigate individual species responses over eight weeks, with an emphasis on the quality of leachate outputs from decomposing litter (from Empetrum hermaphroditum, Vaccinium vitis-idaea, Vaccinium uliginosum). In the assay bi-weekly collections of leachate were analysed for phenol oxidase activity, together with total phenolics and dissolved organic C (DOC). At the end of the assay litter mass loss and respired C were also determined. The initial assessment on field mixed-community litter found no enhanced UV-B treatment (henceforth: ‘UV-B treatment’) effect on phenol oxidase activity. However, in the controlled laboratory mesocosm assay, significant species-specific effects of the UV-B treatment were evident, with increased phenol oxidase activity in V. vitis-idaea leachate (P < 0.001) and a significant reduction (P = 0.05) in respired C. Leachate DOC release from the UV-B treatment was greater in both Vaccinium species and the effect on V. uliginosum was significant (P < 0.05). The UV-B treatment had no effect on the total phenolic concentration of litter or leachates for any species, but there were significant differences in leachate total phenolics, both over time and between species. Also the initial phenolic concentration in leachates from the decomposing litter of E. hermaphroditum was greater than both Vaccinum species. Results suggest a species specific role for UV-B in influencing enzyme function and decomposition, dependent on individual traits. This has implications for decomposition dynamics in this system and more widely. Our study highlights the value of using a laboratory assay to gain a mechanistic understanding the species level impacts of a global change factor (UV-B) on decomposition, which are otherwise obscured by community-level responses and difficult to determine under field conditions.