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.     

In vitro interactions between ectomycorrhizal fungi and ericaceous plants

In view of the close association between ericaceous shrubs and ectomycorrhizal trees in forest ecosystems, the interaction between ectomycorrhizal basidiomycetes and the hair roots of four typical ericoid mycorrhizal hosts was investigated in vitro. Seedlings of Vaccinium myrtillus, V. vitis-idaea, V. macrocarpon and Calluna vulgaris were inoculated with each of four ectomycorrhizal basidiomycetes from different phylogenetic groups (Laccaria bicolor, Lactarius musteus, Suillus variegatus and Tomentellopsis submollis) in a low carbon and nutrient agar-cellophane culture system. Two ericoid mycorrhizal Helotiales ascomycetes (Meliniomyces bicolor in the Rhizoscyphus ericae aggregate and a mycobiont out of the Rhizoscyphus ericae aggregate) were included for comparison. Interactions between fungi and hair roots ranged from neutral to surface attachment, and the formation of intracellular hyphal coils. Root and shoot responses to inoculation were different between the host/fungus combinations. The ectomycorrhizal fungus L. bicolor formed extensive intracellular colonization, spreading cell-to-cell with multiple hyphal entry points and intracellular hyphal coils with single entry points in C. vulgaris and V. macrocarpon epidermal cells respectively, however, no significant effects on plant growth were detected. Meliniomyces bicolor formed intracellular hyphal coils in the epidermal cells of V. myrtillus and V. macrocarpon but not the other host spp. The M. bicolor isolate stimulate V. myrtillus root length about 2.5 times. Interestingly, although the unknown ascomycete strain out of the Rhizoscyphus ericae aggregate formed intracellular hyphal coils in epidermal cells of all host plants, it suppressed the growth of C. vulgaris, V. myrtillus, and V. vitis-idaea but not to V. macrocarpon. Further and more detailed experimentation under more ecological realistic conditions for a longer period of time is needed.     

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.     

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.     

Comparative ecophysiology of two representativeQuercus species appearing in different stages of succession

Ecophysiological comparisons were made of the growth and photosynthetic characteristics between seedlings of deciduousQuercus serrata and evergreenQuercus myrsinaefolia. Q. myrsinaefolia seedlings naturally occurring in secondary coppice forests showed exponential-like growth in height with age, while sympatricQ. serrata seedlings were considerably smaller in height, their growth being limited by shortage of light. The photosynthetic characteristics measured under laboratory conditions showed no bases for the differences in growth between the two species on the forest floor: Light compensation points of the seedlings raised under 5% daylight were almost identical for the two species, being about 6.0 μE·m⁻²·s⁻¹. Growth analysis of seedlings planted in a coppice forest showed that bothQ. serrata andQ. myrsinaefolia could hardly grow during the summer under the shrub layer, when relative photon flux density (RPFD) was 0.9±0.5%. In the winter, when RPFD under the leafless canopy increased to 29.3±2.7%, the dry matter production of the evergreen seedlings ofQ. myrsinaefolia was much improved. Current-year seedlings of the species showed NAR of 0.102±0.021 g·dm⁻²·mo⁻¹ during the winter. Temperature dependency of photosynthesis and increment of leaf temperature by direct solar beam also indicated active photosynthesis ofQ. myrsinaefolia on the forest floor during the winter.     

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.     

Growth and reproduction of dwarf shrubs, Vaccinium myrtillus and V. vitis-idaea, in a severely polluted area

Populations of both Vaccinium myrtillus and V. vitis-idaea in heavily polluted areas around the large nickel–copper smelter at Monchegorsk (NW Russia) persist in small patches scattered over the bare soils. Two non-specific stress indices, leaf fluctuating asymmetry and chlorophyll fluorescence, in plants growing in the most polluted sites were at the same level as in plants growing in unpolluted habitats. Mean mass of a vegetative shoot and current year production of vegetative mass were also similar in polluted and unpolluted sites. Branching in polluted habitats was higher in both species. Annual production of new ramets in V. myrtillus, but not in V. vitis-idaea, was lower in polluted habitats. In both species, proportion of sexually reproducing patches was higher and these patches had more generative shoots and higher fruit yield under pollution impact; the reproductive effort increased with pollution in V. myrtillus but not in V. vitis-idaea. Somatic cost of reproduction in both species was higher in polluted sites, presumably due to both more intensive sexual reproduction and resource limitation in low-nutritive contaminated soil. In both species fruit yield was slightly lower under tree canopies than in gaps. However, in industrial barrens fruit yield was higher than in forests (both in gaps and under the canopies), indicating that enhancement of sexual reproduction in polluted sites cannot be explained by higher light availability alone. Increased allocation to sexual reproduction in heavily polluted habitats, where seedling establishment is hampered by soil toxicity, may indicate the plant strategy to disperse from unfavourable environments.

Zusammenfassung

Sowohl Populationen von Vaccinium myrtillus als auch V. vitis-idaea bleiben in den stark verschmutzten Gebieten rund um die große Nickel-Kupfer-Metallhütte von Monchegorsk (NW Russland) in kleinen Flecken verstreut über den kahlen Boden erhalten. Zwei unspezifische Stressindikatoren, die Blattfluktationsasymmetrie und die Chlorophyllfluoreszenz, waren bei Pflanzen, die in den am stärksten verschmutzten Gebieten wuchsen, auf dem gleichen Niveau wie bei Pflanzen, die in unverschmutzten Gebieten wuchsen. Die mittlere Masse der Vegetationstriebe und die diesjährige Produktion vegetativer Masse waren ebenfalls ähnlich in verschmutzten und unverschmutzten Gebieten. Die Verzweigung war bei beiden Arten stärker in den verschmutzten Gebieten. Die jährliche Produktion von neuen Rameten war bei V. myrtillus im Gegensatz zu V. vitis-idaea geringer in den verschmutzten Gebieten. Bei beiden Arten war der Anteil der sich sexuell reproduzierenden Flecken größer und diese Flecken hatten mehr generative Triebe und höhere Fruchtausbeute unter dem Einfluss von Verschmutzung. Der reproduktive Aufwand nahm bei V. myrtillus im Gegensatz zu V. vitis-idaea zu. Die somatischen Kosten der Reproduktion waren bei beiden Arten in den verschmutzten Gebieten höher, vermutlich sowohl aufgrund der intensiveren sexuellen Reproduktion, als auch aufgrund der Ressourcenlimitierung in den nährstoffarmen, kontaminierten Böden. Bei beiden Arten war die Fruchtausbeute geringfügig kleiner unter Baumkronen als in Lücken. In den industriellen Ödländern war jedoch die Fruchtausbeute größer als in den Wäldern (sowohl in den Lücken als auch unter Baumkronen) und wies darauf hin, dass die Erhöhung der sexuellen Reproduktion in den verschmutzten Gebieten nicht allein durch die höhere Verfügbarkeit von Licht erklärt werden kann. Die erhöhte Allokation zu sexueller Reproduktion in den stark verschmutzen Habitaten, wo die Keimlingetablierung aufgrund der Toxizität der Böden erschwert ist, könnte darauf hinweisen, dass die Strategie der Pflanze ist, sich von ungünstigen Umwelten weg auszubreiten.     

Transpiration in response to variation in microclimate and soil moisture in southeastern deciduous forests

Responses of forests to changes in environmental conditions reflect the integrated behavior of their constituent species. We investigated sap flux-scaled transpiration responses of two species prevalent in upland eastern hardwood forests, Quercus alba in the upper canopy and Acer rubrum in the low to mid canopy, to changes in photosynthetically active radiation above the canopy (Q_o), vapor pressure deficit within the canopy (D), and soil moisture depletion during an entire growing season. Water loss before bud break (presumably through the bark) increased linearly with D, reaching 8% of daily stand transpiration (E_C) as measured when leaf area index was at maximum, and accounting for 5% of annual water loss. After leaves were completely expanded and when soil moisture was high, sap flux-scaled daily E_C increased linearly with the daily sum of Q_o. Species differences in this response were observed. Q. alba reached a maximum transpiration at low Q_o, while A. rubrum showed increasing transpiration with Q_o at all light levels. Daily E_C increased in response to daily average D, with an asymptotic response due to the behavior of Q. alba. Transpiration of A. rubrum showed a greater response to soil moisture depletion than did that of Q. alba. When evaluated at a half-hourly scale under high Q_o, mean canopy stomatal conductance (G_S) of individuals decreased with D. The sensitivity of G_S to D was greater in species with higher intrinsic G_S. Regardless of position in the canopy, diffuse-porous species in this and an additional, more mesic stand showed higher G_S and greater stomatal sensitivity to environmental variation than do ring-porous species.     

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.     

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.     

Growth analysis ofQuercus serrata seedlings withinMiscanthus sinensis grass canopies differing in light availability

The effects of different light regimes on the survival, growth and morphology ofQuercus serrata seedlings were studied in canopies ofMiscanthus sinensis. The seedlings of various ages (0–3 yr) were grown in three light regimes: under a denseM. sinensis canopy (TG plot) receiving 2.5%–8.7% of full sunlight, under a relatively sparse canopy (SG plot) receiving 3.8%–16.1% of light and in an adjacent open site (NG plot). There was a little difference in the survival ofQ. serrata seedlings among the three plots. Height and diameter of stem and total leaf area of the seedlings were significantly lower in the shadier plots. However, the first (bottom) flush of the stem was significantly longer in the TG plot than in the NG and SG plots. Total dry weights of individual 1- and 2-yr-oldQ. serrata seedlings in the TG plot were reduced to about one-twelfth of those in the NG plot. Although the relative proportion in dry weight of each organ did not differ significantly among the plots, leaf area ratio, specific leaf area and stem height per unit dry weight were significantly higher in shadier plots. The leaf area per unit stem height was increased considerably in the sunnier plots.     

Influence of elevated CO2 on canopy development and red:far-red ratios in two-storied stands ofRicinus communis

Vertical structure of plant stands and canopies may change under conditions of elevated CO₂ due to differential responses of overstory and understory plants or plant parts. In the long term, seedling recruitment, competition, and thus population or community structure may be affected. Aside from the possible differential direct effects of elevated CO₂ on photosynthesis and growth, both the quantity and quality of the light below the overstory canopy could be indirectly affected by CO₂-induced changes in overstory leaf area index (LAI) and/or changes in overstory leaf quality. In order to explore such possible interactions, we compared canopy leaf area development, canopy light extinction and the quality of light beneath overstory leaves of two-storied monospecific stands ofRicinus communis exposed to ambient (340 μl l⁻¹) and elevated (610 μl l⁻¹) CO₂. Plants in each stand were grown in a common soil as closed “artificial ecosystems” with a ground area of 6.7 m². LAI of overstory plants in all ecosystems more than doubled during the experiment but was not different between CO₂ treatments at the end. As a consequence, extinction of photosynthetically active radiation (PAR) was also not altered. However, under elevated CO₂ the red to far-red ratio (R:FR) measured beneath overstory leaves was 10% lower than in ecosystems treated with ambient CO₂. This reduction was associated with increased thickness of palisade layers of overstory leaves and appears to be a plausible explanation for the specific enhancement of stem elongation of understory plants (without a corresponding biomass response) under elevated CO₂. CO₂ enrichment led to increased biomass of overstory plants (mainly stem biomass) but had no effect on understory biomass. The results of this study raise the possibility of an important indirect effect of elevated CO₂ at the stand-level. We suggest that, under elevated CO₂, reductions in the R:FR ratio beneath overstory canopies may affect understory plant development independently of the effects of PAR extinction.     

Relationship between plant type and canopy apparent photosynthesis in maize (Zea mays L.)

Experiments were conducted to study the effect of plant type on canopy photosynthesis under field conditions. A chamber made of aluminium frame covered with clear plastic material was used to estimate canopy CO₂-exchange rates over a land area of 1.33 m². The plant type of maize “Shendan 7” [planophile type, original-type (OT)] was changed to erectophile type [altered-type (AT)] at silking stage. The rates of canopy apparent photosynthesis (CAP) were measured in both types of maize grown at five plant densities during the reproductive phase. It was shown that AT canopies had greater rates (about 17.2%) of CAP than did OT canopies and the yield increased by about 5.9–8.6% in AT canopies. The vertical distribution of photosynthetic photon flux density and CO₂ concentration in AT canopies were more uniform than those in OT ones. It was suggested that the compact architecture of maize canopy was excellent for photosynthesis and yield formation.     

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.     

UV-B and PAR in single and mixed canopies grown under different UV-B exclusions in the field

The purpose of this study was to investigate whether differences in canopy architecture due to the investigated species (planophile versus erectophile, single versus mixed canopies) or to UV-B effects on plant morphology, lead to differences in UV-B and UV-B/PAR doses within canopies.The development of a very small (10 mm diameter) UV-B and PAR sensor on a long 5 mm wide stick allowed us to measure the penetration of UV-B and PAR in single and mixed canopies of the grass Dactylis glomerata and white clover, Trifolium repens. The plants were grown in greenhouses covered with different thicknesses (3 and 5 mm) of UV-transmittant plexi (12 and 18% UV-B exclusion).For clover, a planophile vegetation, radiation penetration was very low for both UV-B and PAR. UV-B penetration was much less than for PAR, resulting in low UV-B/PAR ratio's within the canopy. This is explained by the low UV-B transmittance of the leaves (<0.1 %) in combination with the planophile leaves.In the grass species, both UV-B and PAR penetrated much deeper into the canopy due to the erectophile structure. The difference between UV-B and PAR penetration was generally quite small except in very tall canopies.The mixed species canopies showed results comparable to the clover canopies. Due to the strongly increased grass growth in these plots, light penetration was generally much lower than in the single species cultures. The increased growth of grass in these mixed plots could be linked to the lower UV-B/PAR dose they received.In plots grown under the higher UV-B level there was a relative decrease in UV-B/PAR ratio within the canopy for both species, compared to canopies from the lower UV-B greenhouses. This could not be explained by changes in leaf angle or biomass, but might be linked to the increase in leaf transmittance of PAR.     

The influence of elevated CO2 on non-structural carbohydrate distribution and fructan accumulation in wheat canopies

We grew 2.4 m² wheat canopies in a large growth chamber under high photosynthetic photon flux (1000 μmol m⁻² s⁻¹) and using two CO₂ concentrations, 360 and 1200 μmol mol⁻¹. Photosynthetically active radiation (400–700 nm) was attenuated slightly faster through canopies grown in 360μmol mol⁻¹ than through canopies grown in 1200μmol mol⁻¹, even though high-CO₂ canopies attained larger leaf area indices. Tissue fractions were sampled from each 5-cm layer of the canopies. Leaf tissue sampled from the tops of canopies grown in 1200μmol mol⁻¹ accumulated significantly more total non-structural carbohydrate, starch, fructan, sucrose, and glucose (p≤ 0.05) than for canopies grown in 360μmol mol⁻¹. Non-structural carbohydrate did not significantly increase in the lower canopy layers of the elevated CO₂ treatment. Elevated CO₂ induced fructan synthesis in all leaf tissue fractions, but fructan formation was greatest in the uppermost leaf area. A moderate temperature reduction of 10 °C over 5d increased starch, fructan and glucose levels in canopies grown in 1200μmol mol⁻¹, but concentrations of sucrose and fructose decreased slightly or remained unchanged. Those results may correspond with the use of fructosyl-residues and release of glucose when sucrose is consumed in fructan synthesis.     

Early-inbreeding depression inVaccinium myrtillus andV. vitis-idaea

Summary Fertility parameters were estimated inVaccinium myrtillus andV. vitis-idaea after self- and cross-pollinations performed in growth chamber. We showed a drastic decrease in fertility after self-pollination as compared to cross-pollination. Number of plump seeds per berry was compared with previous field data. In both species, growth room conditions improved plump seed number after cross-pollination but not after self-pollination. In addition, in order to enhance resources supply to young developing fruits, cytokinin application was tested inV. myrtillus but no effect was detected. We hypothesize that the partial self-sterility is due to inbreeding depression based on the expression of recessive lethals.     

Photomodulation of axis extension in sparse canopies : role of the stem in the perception of light-quality signals of stand density

A fiber optic probe inserted into plant tissues was used to investigate the effects of canopy density on the light environment in different organs. The red:far-red ratio inside the stem of Datura ferox L. seedlings and the estimated phytochrome photoequilibrium were strongly reduced by the presence of neighbors forming canopies too sparse to cause any mutual shading at the level of the leaves. In such canopies, changes in plant density had little effects on the light regime inside the leaves of the succulent Aeonium haworthii (S.D.) Webb et Berth., particularly when the lamina was kept nearly normal to the direct rays of the sun. In field experiments using D. ferox and Sinapis alba L. seedlings, the elongation of the internodes responded to various types of localized light-quality treatments that simulated different plant densities in sparse canopies. The responses were quantitatively similar to those elicited by changes in plant density. The evidence supports the hypothesis that, in stands formed by plants of similar size, the red:far-red ratio of the light that impinges laterally on the stems is among the earliest environmental cues that allow plants to detect local canopy density and adjust axis extension accordingly.     

The relative importance of allelopathy in interference: the effects of an invasive weed on a native bunchgrass

The relative importance of allelopathy and resource competition in plant-plant interactions has been vigorously debated but seldom tested. We used activated carbon to manipulate the effects of root exudates of Centaurea maculosa, a noxious weed in much of western North America, on root elongation rates and growth of the native bunchgrass Festuca idahoensis in order to investigate the relative importance of allelopathy in the total interference of Centaurea. In root observation chambers, Festuca root elongation rates decreased to ᅢ% of the control, beginning 4 days before contacting Centaurea roots in silica sand. However, when activated carbon, which has a high affinity for adsorbing to organic compounds, was added to the sand the effects of Centaurea roots on Festuca root elongation were reduced. In other experiments, Festuca plants were 50% smaller when grown with Centaurea than with conspecifics in pure silica sand. However, Festuca grown with Centaurea in mixtures of sand and activated carbon were 85% larger than Festuca grown with Centaurea in silica sand without carbon. These results suggest that allelopathy accounts for a substantial proportion of the total interference of Centaurea on Festuca, shifting the balance of competition in favor of Centaurea. However, Centaurea outperformed Festuca even in the presence of activated carbon, demonstrating the importance of the combined roles of resource competition and allelopathy.