Rooting strategy of naturally regenerated beech in Silver birch and Scots pine woodlands

This study investigated the belowground development and strategy of late-successional European beech (Fagus sylvatica L.) in ageing natural Scots pine (Pinus sylvestris L.) and Silver birch (Betula pendula Roth.) woodlands in a French volcanic mid-elevation area. For this purpose root biomass, root profile and fine-root architecture of competitor trees were examined in 53 mixed pine–beech and 42 birch–beech woodlands along a stand maturation gradient, using the root auger technique (0–75-cm). The total beech fine-root biomass highly correlated with aerial dimensions such as stem height and girth, whereas it moderately correlated with its age, thus indicating the effects of competition. Basic stand biometric data such as stand density and basal area had no significant effect on beech root biomass. Conversely, competition indices taking into account the vertical dimensions of competitor trees were efficient, probably due to redundancy with beech height. At similar age and height, beeches under birch had a greater belowground development than beeches under pine. Each species exhibited specific rooting pattern and plasticity of fine-root architecture along the gradients of stand maturation and competition. Beech had a heart-shaped rooting habit in both mixings, which strongly increased along stand maturation. Its fine-root system adopted a foraging strategy to respond to increasing stand competition. The Scots pine fine-root system was plate-like and showed a low morphological plasticity, thus presumably a conservative strategy. Silver birch exhibited a high biomass and a foraging capacity in the topsoil but a loose root system in the subsoil. The coexistence of pine and beech roots in the upper soil presumably leads to a high belowground competition. Beech root system becomes predominant throughout the soil profile and it adopts an efficient foraging strategy, but at the expense of its belowground development. Conversely, the niche partitioning strategy between beech and birch may explain why beech develops strongly belowground in spite of the fact that birch has a dense rooting and a competitive fine-root architecture. As a consequence, beech mid-term regeneration and development may be facilitated under birch as compared with pine.     

两种落叶分解过程中养分和碳素的行为

利用分解袋法对日本亚高山针叶林的针叶(Abies veitchii Lindl,and A.mariesi Mast.)和阔叶(Betula ermanii Cham.and B.corylifolia Regal.et Maxim．)凋落物进行了分解实验研究。结果表明尽管分解初期的两种凋落物的养分以及分解后期凋落物剩余重量差异很大,但两种凋落物养分浓度在分解后期(30个月以后)趋于一致。这种趋同现象在不同养分中有不同的趋同机制。氮元素浓度升高到分解后期浓度差变小,这种现象是由于分解菌的固持作用及受木质素的束缚所致;钾和镁在分解初期浓度急剧下降,进而浓度差变小,是由于淋溶作用所致。在分解过程中这些元素非常容易被淋溶掉,直到和土壤中的浓度达到一致为止。钙是结构元素,它的行踪和有机物组分有密切关系。由于分解后期有机组分木质化和腐殖质化进而浓度趋同,所以钙的浓度也相应趋同。     

Seed bank spatial pattern in a temperate secondary forest

Abstract. Seed bank spatial pattern was studied in a secondary forest dominated by Fagus sylvatica and Betula celtiberica in the Urkiola Natural Park (N Spain). Soil samples were taken every 2 m in a regular grid (196 points) and divided into two fractions (0–3 cm and 3–10 cm deep). The viable seed bank was studied by monitoring seedling emergence for ten months. The effect of different factors on seed bank composition and patterning was analysed using constrained ordination as a hypothesis testing tool. Furthermore, the existence of spatial autocorrelation was evaluated by geostatistical analysis. Seed density was high, 7057 seed.m^?2, with a few species dominating. Species composition in the various layers were significantly correlated. The seed bank showed significant spatial structure, which was partially explainable by the spatial structure of the canopy and understorey vegetation. Spatial clumping from 0–8 m was observed in seed bank density and composition, mainly due to the pattern of two abundant taxa Juncus effusus and Ericaceae. The Ericaceae seed bank was related to the spatial distribution of dead stumps of Erica arborea. J. effusus was not present in the above‐ground vegetation, which indicates that its seed bank was formed in the past. As expected, the seed bank of this forest reflects its history, which is characterized by complex man‐induced perturbations. The seed bank appears to be structured as a consequence of contrasting driving forces such as canopy structure, understorey composition and structural and microhabitat features.     

Stand structure and regeneration in a Kamchatka mixed boreal forest

Abstract. A 1‐ha plot was established in a Betula platyphylla‐Picea ajanensis mixed boreal forest in the central Kamchatka peninsula in Russia to investigate stand structure and regeneration. This forest was relatively sparse; total density and stand basal area were 1071/ha and 25.8 m²/ha, respectively, for trees > 2.0 cm in trunk diameter at breast height (DBH). 25% of Betula regenerated by sprouting, and its frequency distribution of DBH had a reverse J‐shaped pattern. In contrast, Picea had a bimodal distribution. The growth rates of both species were high, reaching 20 m in ca. 120 yr. The two species had clumped distributions, especially for saplings. Betula saplings were not distributed in canopy gaps. Small Picea saplings were distributed irrespective of the presence/absence of gaps, while larger saplings aggregated in gaps. At the examined spatial scales (6.25–400 m²) the spatial distribution of Betula saplings was positively correlated with living Betula canopy trees and negatively with dead Picea canopy trees. This suggests that Betula saplings regenerated under the crowns of Betula canopy trees and did not invade the gaps created by Picea canopy trees. The spatial distribution of Picea saplings was negatively correlated with living and dead Betula canopy trees and positively with dead Picea canopy trees. Most small Picea seedlings were distributed under the crowns of Picea trees but not under the crowns of Betula trees or in gaps. This suggests that Picea seedlings establish under the crowns of Picea canopy trees and can grow to large sizes after the death of overhead Picea canopy trees. Evidence of competitive exclusion between the two species was not found. At a 20 m × 20 m scale both skewness and the coefficient of variation of DBH frequency distribution of Picea decreased with an increase in total basal area of Picea while those of Betula were unchanged irrespective of the increase in total basal area of Betula. This indicates that the size structure of Picea is more variable with stand development than that of Betula on a small scale. This study suggests that Betula regenerates continuously by sprouting and Picea regenerates discontinuously after gap formation and that the species do not exclude each other.     

The effects of long-term elevated UV-B on the growth and phenolics of field-grown silver birch (Betula pendula)

The effects of long‐term elevated UV‐B radiation on silver birch (Betula pendula Roth) seedlings were studied over three growing seasons in an outdoor experiment in Finland started 64 days after germination. One group of seedlings was exposed to a constant 50% increase in UV‐B_CIE radiation, which corresponds to 20–25% of ozone depletion; another group received a small increase in UV‐A radiation and a third (the control group) received ambient solar radiation. Changes in growth appeared during the third growing season; the stems of the UV‐B treated seedlings were thinner and their height tended to be shorter compared with that of the control seedlings. In contrast, there were no UV‐B effects on biomass, bud burst, bud dry weights, leaf area, rust frequency index or chlorophyll concentrations in any of the summers. During the three‐year study, the flavonols were significantly increased by the elevated UV‐B only in the first growing season. The responses varied greatly among individual compounds; the most induced were the quercetin glycosides, while the main flavonols, myricetins, were reduced by the UV‐A control treatment. In the second summer phenolic acids, such as 3,4′‐dihydroxypropiophenone‐3‐glucoside, neochlorogenic acid and 5‐coumarylquinic acid, were increased by the UV‐B treatment. In the third year, the constitutive concentrations of phenolics were not affected by the UV‐B treatment.     

Effects of isolates of ectomycorrhizal fungi and endophytic Mycelium radicis atrovirens that were dominant in soil from disturbed sites on growth of Betula platyphylla var. japonica seedlings

Effects of ectomycorrhizal fungi and endophytic Mycelium radicis atrovirens Melin (MRA) on growth of Betula platyphylla var. japonica seedlings were investigated under aseptic culture conditions. Three isolates of ectomycorrhizal fungi and two isolates of MRA were used. One MRA isolate was Phialocephala fortinii. Previous field work revealed that these isolates were dominant on the roots of B. platyphylla var. japonica seedlings grown in a mineral subsoil that had been exposed by the removal of surface soil. After a 100-day incubation, the growth of the seedlings was significantly enhanced by the colonization of these ectomycorrhizal fungal isolates as compared with uninoculated seedlings. In contrast, the growth of seedlings was retarded by the colonization of the MRA isolates. The growth of seedlings that were co-inoculated with ectomycorrhizal fungi and MRA was similar to that of uninoculated seedlings in most cases. These results suggest that ectomycorrhizal fungi have a beneficial effect on the growth of B. platyphylla var. japonica seedlings and that they suppress the deleterious effect of MRA. Thus, these ectomycorrhizal fungi probably have an important role in establishing B. platyphylla var. japonica seedlings during the initial stage of re-vegetation following site disturbance by the removal of surface soil.     

A computer-operated routine of gas exchange and optical measurements to diagnose photosynthetic apparatus in leaves

Photosynthesis is a complex process whose rate is affected by many biochemical and biophysical factors. Fortunately, it is possible to determine, or at least estimate, many of the most important parameters using a combination of optical methods and gas transient analyses. We describe here a computer‐operated routine that has been developed to make detailed assessments of photosynthesis at a comprehensive level. The routine comprised the following measurements: steady‐state light and CO₂ response curves of net CO₂ assimilation at 21 and 2 kPa O₂; transients from limiting to different saturating CO₂ concentrations at 2 kPa O₂; post‐illumination CO₂ fixation transient; dark–light induction of O₂ evolution; O₂ yield from one saturating single‐turnover flash; chlorophyll fluorescence F₀, F_s and F_m during the light and CO₂ response curves; leaf transmission at 820 nm (P700⁺) during the light and CO₂ response curves; post‐illumination re‐reduction time of P700⁺. The routine was executed on a two‐channel fast‐response gas exchange measurement system (A. Laisk and V. Oja: Dynamic Gas Exchange of Leaf Photosynthesis. CSIRO, Canberra, Australia). Thirty‐six intrinsic characteristics of the photosynthetic machinery were derived, including quantum yield of CO₂ fixation (Y_CO2), time constant of P700 re‐reduction (τ′), relative optical cross‐sections of PSII and PSI antennae (a_II, a_I), PSII and PSI density per leaf area unit, plastoquinone pool, total mesophyll resistance, mesophyll diffusion resistance, V_m, K_m(CO₂) and CO₂/O₂ specificity of Rubisco, RuBP pool at CO₂ limitation (assimilatory charge). An example of the routine and calculations are shown for one leaf and data are presented for leaves of 8‐year‐old‐trees of two birch clones growing in Suonenjoki Forest Research Station, Finland, during summer 2000. Parameters Y_CO2, basic τ′, a_II, a_I, K_m(CO₂) and K_s varied little in different leaves [relative standard deviation (RSD) < 7%], other parameters scattered widely (RSD typically 10–40%). It is concluded that the little scattered parameters are determined by basic physico‐chemical properties of the photosynthetic machinery whereas the widely scattered parameters are adjusting to growth conditions. The proposed non‐destructive routine is suitable for diagnosing the photosynthetic machinery of leaves and may be applied in plant ecophysiology and in genetic engineering of plants.     

Root discoloration and shoot symptoms in silver birch after Phytophthora infection in vitro

• There are no records of established plant pathogenic Phytophthora species in Finnish forests, but they are likely in the future. Therefore, the effects of Phytophthora inoculations on young, ca. 2‐month‐old silver birch (Betula pendula) seedling roots and shoots were investigated.
• Visual inspection of dark discoloration, direct PCR and re‐isolation, and detailed root morphology analyses were used to evaluate the effects of Phytophthora inoculation on roots. Symptoms in leaves and stems were also recorded.
• Phytophthora was successfully re‐isolated from 67% of the surface‐sterilized roots of inoculated seedlings, but not from the non‐inoculated control seedlings. Dark discolorations were found more often in the root segments of inoculated seedlings than in control seedlings. In the Phytophthora‐treated seedlings, discoloured root segments were usually linked and found primarily in the main root or lateral roots attached to it, whereas in the control seedlings a few single discoloured root segments were scattered throughout the root systems. The number of root segments was lower in the inoculated than in the control seedlings, indicating root loss after Phytophthora inoculation. In the shoots of inoculated birches, leaf and shoot wilting was observed.
• The appearance of wilting in shoots without visible dark discoloration in the base of stems indicated that symptoms originated from roots inoculated with Phytophthora.

     

Early post-fire tree regeneration in a Picea-Vaccinium forest in northern Sweden

Abstract. Age and size structure of saplings of Picea abies, Pinus sylvestris and Betula pubescens were examined in a 26-yr old forest fire area in a Picea abies-Vaccinium myrtillus forest in northern Sweden. Picea, which is a shade-tolerant species, had its maximum regeneration prior to the shade-intolerant Pinus. The shift from Picea to Pinus regeneration in the late 1970s, did not seem to be related to variations in summer temperature. Instead, it is suggested that Picea established in the shade created by dead trees, and, that increased reindeer browsing of Betula in combination with a simultaneous thinning of the tree layer, favoured Pinus recruitment. These regeneration patterns do not confirm conventional views of post-fire succession in Sweden.