Competition increasingly dominates the responsiveness of juvenile beech and spruce to elevated CO2 and/or O3 concentrations throughout two subsequent growing seasons

Saplings of Fagus sylvatica and Picea abies were grown in mono‐ and mixed cultures in a 2‐year phytotron study under all four combinations of ambient and elevated ozone (O₃) and carbon dioxide (CO₂) concentrations. The hypotheses tested were (1) that the competitiveness of beech rather than spruce is negatively affected by the exposure to enhanced O₃ concentrations, (2) spruce benefits from the increase of resource availability (elevated CO₂) in the mixed culture and (3) that the responsiveness of plants to CO₂ and O₃ depends on the type of competition (i.e. intra vs. interspecific). Beech displayed a competitive disadvantage when growing in mixture with spruce: after two growing seasons under interspecific competition, beech showed significant reductions in leaf gas exchange, biomass development and crown volume as compared with beech plants growing in monoculture. In competition with spruce, beech appeared to be nitrogen (N)‐limited, whereas spruce tended to benefit in terms of its plant N status. The responsiveness of the juvenile trees to the atmospheric treatments differed between species and was dominated by the type of competition: spruce growth benefited from elevated CO₂ concentrations, while beech growth suffered from the enhanced O₃ regime. In general, interspecific competition enhanced these atmospheric treatment effects, supporting our hypotheses. Significant differences in root : shoot biomass ratio between the type of competition under both elevated O₃ and CO₂ were not caused by readjustments of biomass partitioning, but were dependent on tree size. Our study stresses that competition is an important factor driving plant development, and suggests that the knowledge about responses of plants to elevated CO₂ and/or O₃, acquired from plants growing in monoculture, may not be transferred to plants grown under interspecific competition as typically found in the field.     

Above-ground space sequestration determines competitive success in juvenile beech and spruce trees

A 2-yr phytotron study was conducted to investigate the intra- and inter-specific competitive behaviour of juvenile beech (Fagus sylvatica) and spruce (Picea abies). Competitiveness was analysed by quantifying the resource budgets that occur along structures and within occupied space of relevance for competitive interaction. Ambient and elevated CO(2) and ozone (O(3)) regimes were applied throughout two growing seasons as stressors for provoking changes in resource budgets, growth and allocation to facilitate the competition analysis. The hypothesis tested was that the ability to sequester space at low structural cost will determine the competitive success. Spruce was a stronger competitor than beech, as displayed by its higher above-ground biomass increments in mixed culture compared with monoculture. A crucial factor in the competitive success of spruce was its ability to enlarge crown volume at low structural costs, supporting the hypothesis. Interspecific competition with spruce resulted in a size-independent readjustment of above-ground allocation in beech (reduced leaf : shoot biomass ratio). The efficient use of resources for above-ground space sequestration proved to be a parameter that quantitatively reflects competitiveness.     

A new scenario for the quaternary history of European beech populations: palaeobotanical evidence and genetic consequences

Here, palaeobotanical and genetic data for common beech (Fagus sylvatica) in Europe are used to evaluate the genetic consequences of long-term survival in refuge areas and postglacial spread. Four large datasets are presented, including over 400 fossil-pollen sites, 80 plant-macrofossil sites, and 450 and 600 modern beech populations for chloroplast and nuclear markers, respectively. The largely complementary palaeobotanical and genetic data indicate that: (i) beech survived the last glacial period in multiple refuge areas; (ii) the central European refugia were separated from the Mediterranean refugia; (iii) the Mediterranean refuges did not contribute to the colonization of central and northern Europe; (iv) some populations expanded considerably during the postglacial period, while others experienced only a limited expansion; (v) the mountain chains were not geographical barriers for beech but rather facilitated its diffusion; and (vi) the modern genetic diversity was shaped over multiple glacial-interglacial cycles. This scenario differs from many recent treatments of tree phylogeography in Europe that largely focus on the last ice age and the postglacial period to interpret genetic structure and argue that the southern peninsulas (Iberian, Italian and Balkan) were the main source areas for trees in central and northern Europe.     

Climate change reverses the competitive balance of ash and beech seedlings under simulated forest conditions

This study identifies the important role of climate change and photosynthetic photon flux density (PPFD) in the regenerative competence of ash and beech seedlings in 12 inter- and intra-specific competition designs in simulated mixed ash-beech forest gaps under conditions of non-limiting soil volume, water and nutrient supply. The growth conditions simulated natural forest conditions as closely as possible. Simulations were performed by growing interacting seedling canopies for one season in temperature-regulated closed-top chambers (CTCs). Eight CTCs were used in a factorial design with replicate treatments of [CO2] x temperature x PPFD x competition design. [CO2] tracked ambient levels or was 360 micromol mol-1 higher. Temperature tracked ambient levels or was 2.8 degrees C higher. PPFD on two plant tables inside each CTC was 16% and 5% of open-field levels, respectively, representative of typical light flux levels in a natural forest gap. In several of the competition designs, climate change made the ash seedlings grow taller than the beech seedlings and, at the same time, attain a larger leaf area and a larger total biomass. Advantages of this type for ash were found particularly at lower PPFD. There was a positive synergistic interaction of elevated temperature x [CO2] for both species, but more so for ash. There are many uncertainties when a study of chambered seedlings is to be projected to real changes in natural forests. Nevertheless, this study supports a possible future shift towards ash in north European, unmanaged, mixed ash-beech forests in response to the predicted climate change.     

Carbon and nitrogen dynamics in acid detergent fibre lignins of beech (Fagus sylvatica L.) during the growth phase

To study the incorporation of carbon and nitrogen in different plant fractions, 3‐year‐old‐beech (Fagus sylvatica L.) seedlings were exposed in microcosms to a dual‐labelling experiment employing ¹³C and ¹⁵N throughout one season. Leaves, stems, coarse and fine roots were harvested 6, 12 and 18 weeks after bud break (June to September) and used to isolate acid‐detergent fibre lignins (ADF lignin) for the determination of carbon and nitrogen and their isotope ratios. Lignin concentrations were also determined with the thioglycolic acid method. The highest lignin concentrations were found in fine roots. ADF lignins of all tissues analysed, especially those of leaves, also contained significant concentrations of nitrogen. This suggests that lignin‐bound proteins constitute an important cell wall fraction and shows that the ADF method is not suitable to determine genuine lignin. ADF lignin should be re‐named as ligno‐protein fraction. Whole‐leaf biomass was composed of 50 to 70% newly assimilated carbon and about 7% newly assimilated nitrogen; net changes in the isotope ratios were not observed during the experimental period. In the other tissues analysed, the fraction of new carbon and nitrogen was initially low and increased significantly during the time‐course of the experiment, whereas the total tissue concentrations of carbon remained almost unaffected and nitrogen declined. At the end of the experiment, the whole‐tissue biomass and ADF lignins of fine roots contained about 65 and 50% new carbon and about 50 and 40% new nitrogen, respectively. These results indicate that significant metabolic activity was related to the formation of structural biopolymers after leaf growth, especially below‐ground and that this activity also led to a substantial binding of nitrogen to structural compounds.     

黑莓对水分、盐分和低温逆境的适应性

在南京盆栽条件下,黑莓（Rubus L.spp.)“Hull”品种1年生扦插苗在土壤相对含水量为70%以上时,地上部分和地下部分均生长良好,正常开花结实;土壤相对含水量下降到50%-60%时,地上部分生长明显减弱,地下部分生长受到一定影响,但植株都能存活;土壤相对含水量下降到45%-50%时,地上部分及地下部分均生长不良,部分植株死亡;土壤相对含水量降到30%-35%时,植株无法生存。黑莓“Hull”品种扦插苗是对盐（NaCl)敏感的植物,中度（0.2%)以上盐浓度对生长有明显的抑制作用,相对于地上部分和根系,根颈部所受的影响较小,表现为地上部无新的枝条萌出,无叶片产生;地下部无新根产生;根颈部萌芽数量减少,长度变小,随着盐浓度的增加,生长减慢甚至死亡。轻度（0.1%)盐处理未见有抑制作用。黑莓不同品种的枝条抗冻性不同,所测定的3个品种中,“HuLL”抗冻性最差,LT50为-25.5℃,“Chester”与“Navoho”的抗冻性相近,LT50均为-29℃。     

Plant-mediated nitrous oxide emissions from beech (Fagus sylvatica) leaves

Nitrous oxide (N2O) emission estimates from forest ecosystems are based currently on emission measurements using soil enclosures. Such enclosures exclude emissions via tall plants and trees and may therefore underestimate the whole-ecosystem N2O emissions. Here, we measured plant-mediated N2O emissions from the leaves of potted beech (Fagus sylvatica) seedlings after fertilizing the soil with 15N-labelled ammonium nitrate (15NH4(15)NO3), and after exposing the roots to elevated concentrations of N2O. Ammonium nitrate fertilization induced N2O + 15N2O emissions from beech leaves. Likewise, the foliage emitted N2O after beech roots were exposed to elevated concentrations of N2O. The average N2O emissions from the fertilization and the root exposure experiments were 0.4 and 2.0 microg N m(-2) leaf area h(-1), respectively. Higher than ambient atmospheric concentrations of N2O in the leaves of the forest trees indicate a potential for canopy N2O emissions in the forest. Our experiments demonstrate the existence of a previously overlooked pathway of N2O to the atmosphere in forest ecosystems, and bring about a need to investigate the magnitude of this phenomenon at larger scales.     

世界黑莓产业发展和研究现状及前景

在查阅大量文献资料的基础上,结合作者的研究成果,对国内外黑莓(Rubus spp.)产业发展及研究现状进行了综述,并分析了存在的问题,阐述了中国黑莓产业的发展优势和产业化前景.目前黑莓在全世界的种植面积约为20 035 hm2,年产量1.5×105 ～2.0×105 t,栽培区域遍及各大洲,其中塞尔维亚的黑莓种植面积位居世界第一,美国的黑莓年产量居世界第一,亚洲仅中国种植黑莓;近几年来,罗马尼亚、波兰、墨西哥、智利、匈牙利、中国以及美国的黑莓种植面积增长较快;因黑莓极不易贮存,绝大多数鲜果用于加工.国外以常规育种技术为主、分子技术为辅培育出许多黑莓品种,栽培技术已经非常成熟,成功实现了机械化采收,并逐步推广黑莓的有机生产体系和大棚栽培技术.国外黑莓生产中面临的主要问题是经济、冻害和病虫害等.黑莓于1986年引入中国,从1994年开始推广,至2010年种植面积约4 500 hm2,主要集中在江苏省的溧水县、赣榆县和溧阳市,90％以上鲜果加工成速冻果出口.在国外引种和国内野生种质资源收集的基础上,中国的研究者开展了黑莓的选种和育种研究,并培育和筛选出适宜于本地种植的黑莓优良品种;黑莓的栽培、加工技术研究也取得了显著的成果,并研制出一些加工产品.当前中国黑莓产业主要面临主栽品种单一、农药残留量超标、栽培条件差等方面的问题,制约了黑莓产业的发展.由于黑莓适于低山丘陵栽培,具有结果早、见效快、营养价值高等优势,在中国具有广阔的发展前景.     

Bioclimate and growth history affect beech lifespan in the Italian Alps and Apennines

When site factors reduce growth rates, tree lifespan tends to increase. This study investigates processes leading to such inverse relationship in Fagus sylvatica stands distributed along two elevation gradients, with an emphasis on climatic response, suppression periods, and growth trends. Dendrochronological records from old‐growth beech populations sampled at different elevations within two different bioclimatic regions (Alps vs. Apennines), were used to investigate factors that control tree lifespan. Differences between old‐growth (12) and nearby managed (15) stands were used to assess effects of silvicultural practices on maximum age. Logging reduced tree lifespan not only by removing older trees, but also by reducing the number of years beech individuals spent in the shaded understory. Tree lifespan and growth rates were affected by climate (spring–summer temperature) and were inversely related to one another along elevation gradients. The greatest lifespan was observed in old‐growth high‐mountain populations, and was related not only to slower growth due to a shorter growing season, but also to multidecadal periods of growth suppression during the initial development stages in the understory (i.e., slower growth rates at the youngest cambial ages). Past unfavorable climatic periods (in this case, the Little Ice Age) also helped increase tree lifespan. Using a linear model, we estimated a reduction in beech lifespan of 23 ± 5 years for each degree of warming. Basal area increment of trees with the maximum observed lifespan showed an increasing trend over time. Because growth of old (>300 years) trees has increased in the Alps, while it has recently declined in the Apennines, different bioclimatic regions can have opposite responses to global climatic change. In the next decades, if warming continues, beech lifespan could be reduced in the Alps by faster growth and in the Apennines by drought‐induced mortality.     

Frost and drought: Effects of extreme weather events on stem carbon dynamics in a Mediterranean beech forest

The effects of short-term extreme events on tree functioning and physiology are still rather elusive. European beech is one of the most sensitive species to late frost and water shortage. We investigated the intra-annual C dynamics in stems under such conditions. Wood formation and stem CO₂ efflux were monitored in a Mediterranean beech forest for 3 years (2015–2017), including a late frost (2016) and a summer drought (2017). The late frost reduced radial growth and, consequently, the amount of carbon fixed in the stem biomass by 80%. Stem carbon dioxide efflux in 2016 was reduced by 25%, which can be attributed to the reduction of effluxes due to growth respiration. Counter to our expectations, we found no effects of the 2017 summer drought on radial growth and stem carbon efflux. The studied extreme weather events had various effects on tree growth. Even though late spring frost had a strong impact on beech radial growth in the current year, trees fully recovered in the following growing season, indicating high resilience of beech to this stressful event.     

Phosphate and calcium uptake in beech (Fagus sylvatica) in the presence of aluminium and natural fulvic acids

In the present study we examine the effects of Al on the uptake of Ca²⁺ and H₂PO^-₄ in beech (Fagus sylvatica L.) grown in inorganic nutrient solutions and nutrient solutions supplied with natural fulvic acids (FA). All the solutions used were chemically well characterized. The uptake of Al by roots of intact plants exposed to solutions containing 0, 0.15 or 0.3 mM AlCl₃ for 24 h, was significantly less if FA (300 mg l⁻¹) were also present in the solutions. The Ca²⁺(⁴⁵Ca²⁺) uptake was less affected by Al in solutions supplied with FA than in solutions without FA. There was a strong negative correlation between the Al and Ca²⁺ uptake (r²=0.98). When the Al and Ca²⁺ (⁴⁵Ca²⁺) uptake were plotted as a function of the Al³⁺ activity (or concentration of inorganic mononuclear Al), almost the same response curves were obtained for the -FA and +FA treatments. We conclude that FA-complexed Al was not available for root uptake and therefore could not affect the Ca²⁺ uptake. The competitive effect of Al on the Ca²⁺ uptake was also shown in a 5-week cultivation experiment, where the Ca concentration in shoots decreased at an AlCl₃ concentration of 0.3 mM. The effect of Al on H₂PO⁻₄ uptake was more complex. The P content in roots and shoots was not significantly affected, compared with the control, by cultivation for 5 weeks in a solution supplied with 0.3 mM AlCl₃, despite a reduction of the H₂PO⁻₄ concentration in the nutrient solution to about one-tenth. At this concentration Al obviously had a positive effect on H₂PO⁻₄ uptake. The presence of FA decreased ³²P-phosphate uptake by more than 60% during 24 h, and the addition of 0.15 or 0.3 mM AlCl₃ to these solutions did not alter the uptake of ³²P-phosphate.