Diminished vessel diameter as a possible factor in the decline of European ash (Fraxinus excelsior L.)

? The aim of this work was to examine the anatomy and functioning of secondary xylem in stems in relation to the decline of European ash (Fraxinus excelsior L.).

? We tested the hypothesis that declining trees show changes in the structure of wood, which result in impaired water transport.

? Anatomical analyses were carried out on wood samples (comprising all annual rings formed during the 30 years life of the analysed trees) collected at breast height from the main stem of healthy, weakened and dead ash trees. The width of annual wood increments, the diameter and density of earlywood vessels were measured and the theoretical hydraulic conductivity index through the secondary xylem calculated by application of Hagen-Poisseuille’s formula.

? Anatomical characteristics changed both with the age of trees and in response to unfavorable factors. The largest vessels were observed in healthy trees, which implied that they had the highest hydraulic conductivity index, whereas trees considered to be in decline produce smaller vessels and hence had reduced conductivity.

     

SEM-observations on the structural degradation of epistomatal waxes in Picea abies (L.) Karst.—and its possible role in the “Fichtensterben”

The structural degradation of the epistomatal waxes was studied with the scanning electron microscope in needles of five trees of Picea abies (L.) Karst. differing in their symptoms of decline. The shedding of needles was found to be related to the degree of occlusion of stomates and to occur when about 80 to 90 % of stomates became occluded by a flat and solid wax cover resulting from the structural degradation of the epistomatal wax crystalloids. While this structural degradation stage was not reached before the seventh year in the trees classified as healthy, degradation was dramatically accelerated in the declining trees. The physiological consequences of such structural obstruction could explain many results so far obtained in declining trees.     

Element concentrations and storage in the aboveground biomass of limed and unlimed Norway spruce trees at Höglwald

Twenty years after the application of dolomitic limestone to a well growing, mature Norway spruce stand at Höglwald, Southern Germany an amount of magnesium equal to the input by liming has left the rooting zone. However, 80% of the applied calcium was retained by the ecosystem. A detailed biomass investigation was carried out in order to elaborate whether the stand shows any nutritional benefits and if the aboveground tree biomass represents a sink for calcium and magnesium after liming. Stem wood, bark, branches, twigs, and needles were analysed for the major nutrients. Needles showed higher calcium and magnesium concentrations and significantly lower potassium concentrations on the limed plot. Surprisingly, limed trees had significantly lower magnesium concentrations in the bark but higher concentrations in twigs and needles than unlimed trees. In addition, significantly lower concentrations for potassium, sulphur, and phosphorus were measured in the bark of limed trees, whereas the element concentrations in the wood were rather similar between the two treatments. The low potassium concentrations in bark and needles indicate an impeded potassium uptake after liming presumably due to a calcium potassium antagonism. Magnesium is probably used to compensate the relative lack of potassium. The small differences in the calculated nutrient storage of limed and unlimed trees, however, suggest that the aboveground biomass can be excluded as a relevant sink for the applied calcium and magnesium at the Höglwald site.     

Nitrogen forms in bark, wood and foliage of nitrogen-fertilized Pinus sylvestris

Cycling of soluble non-protein N compounds is thought to be indicative of the N-nutritional status of trees. We determined the major N forms in bark, wood and foliage and estimated the dependence of prevalent N forms on N availability in Pinus sylvestris L. trees from northern Sweden. Trees subjected to severe N limitation and trees that had been fertilized with an average 64 kg N ha(-1) year(-1) for 25 years were analyzed. Bark and wood samples were collected by tangentially cryo-sectioning the trunk into 30-microm thick sections, from the bark to the functional xylem. Soluble amino compounds were extracted from the sections for analysis. Sap samples from twigs were obtained by centrifugation, and bark samples from twigs were obtained by tissue extraction. In both needles and bark, arginine dominated the amino-N pool. Because arginine concentrations in needles increased with N fertilization, arginine dominance of the amino-N pool in needles was higher in N-fertilized trees than in control trees. In bark, N fertilization resulted in a large increase in glutamine concentration, so that glutamine accounted for a larger proportion of the amino-N pool in bark in N-fertilized trees than in control trees. Glutamine dominated the amino-N pool in wood of control trees. Nitrogen fertilization resulted in an increased proportion of arginine in the wood amino-N pool. We conclude that the composition of the amino-N pools in bark, wood and foliage is highly sensitive to N supply. The composition of the amino-N pools can contribute to the regulation of tree N-nutritional status, which is mediated by shoot to root signalling by long-distance transport of amino compounds.     

毛白杨形成层的活动周期及其POD同工酶的变化

北京地区生长的毛白杨 (PopulustomentosaCarr .)在春季芽膨大前 ,形成层带已经进入了恢复活动期。在芽展开后一周形成层进入活动期 ,细胞开始分裂 ,细胞层数也有所增加 ,同时分化出未成熟的韧皮部细胞。未成熟韧皮部细胞的数量在形成层活动初期快速增长 ,随后是一个持续而缓慢的连续过程。木质部的产生比韧皮部要晚 3周 ,未成熟的木质部细胞层数在开始变化不大 ,5月底～ 6月底 ,增长速度明显高于初期 ,随后便急剧下降。成熟木质部细胞的出现比成熟的韧皮部细胞要晚约一个月。与其他研究结果不同 ,毛白杨在4月底到 6月底这段时间 ,形成层细胞层数迅速下降 ,未成熟韧皮部细胞也略有减少 ,而未成熟和成熟的木质部细胞则大量增加。木质部在 9月底停止分化 ,较韧皮部要晚约 2 0d。 9月底形成层进入休眠期后 ,形成层细胞层数基本保持不变 ,直到翌年春天恢复活动。毛白杨形成层区域的过氧化物酶 (POD)同工酶在形成层活动期其活性低、酶带少 ,而恢复活动期和休眠期其活性高且酶带多 ,组织化学反应还表明 ,不同时期POD在不同的细胞和组织中的活性不同。本文对POD同工酶在形成层活动周期中的变化与其组织分化的关系进行了讨论。     

Soluble N compound profiles and concentrations in European beech (Fagus sylvatica L.) are influenced by local climate and thinning

We assessed seasonal changes of total soluble nonprotein nitrogen compounds (TSNN) in adult European beech trees (Fagus sylvatica, L.) growing under different local climate during the growing season immediately following a thinning treatment and 3 years later. In both years, samples of leaves, xylem sap and phloem exudates from beech trees growing in thinned and unthinned (control) stands on a dry, warm SW exposed and a cooler, moist NE exposed site were collected in May, July and September. In May of both years, asparagine (Asn) and glutamine (Gln) were most abundant in leaves and xylem, respectively, whereas arginine (Arg) dominated in the phloem. In July, TSNN concentrations decreased in all tissues and sites, but differences in water availability between aspects were reflected in TSNN concentrations. In September, differences in the increase of Arg concentration in the phloem were related to differences in the onset of senescence between treatments. Thinning treatment increased amino compound concentrations of beech tissues in July on both aspects, particularly at the NE thinned site. It is supposed that, the N balance of adult beech is favoured by both, the thinning treatments as well as the cool-moist climate prevailing at the NE aspect.     

Methyl jasmonate and oxalic acid treatment of Norway spruce: anatomically based defense responses and increased resistance against fungal infection

To study the effect of chemical pretreatment on conifer resistance, 13-year-old Norway spruce (Picea abies (L.) Karst.) trees were treated with methyl jasmonate (MJ) or oxalic acid (OxA) on the outer bark and inoculated with the pathogenic blue-stain fungus Ceratocystis polonica (Siem.) C. Moreau 4 weeks later. Both chemicals significantly reduced symptoms of fungal infection, but MJ was more effective than OxA (51 versus 18% reduction in length of necrotic lesions in the phloem relative to untreated control trees). Anatomical examination of treated stem tissues showed that MJ induced extensive formation of traumatic resin ducts in the xylem and extra polyphenolic parenchyma (PP) cells in the secondary phloem between the cambium and the regular annual PP cell layer. No traumatic resin ducts were formed after treatment with OxA, and the coverage of extra PP cells in OxA-treated tissues was not significantly higher than in the controls. The anatomically based defense reactions induced by MJ were similar to the reactions observed after pathogen infection, mechanical wounding and bark beetle attack. Neither MJ nor OxA had apparent phytotoxic effects on Norway spruce at the concentrations used, with needle and stem tissues of all trees appearing normal without visible symptoms of toxicity. However, trees treated with MJ had 30% less radial sapwood growth than control trees. In conclusion, MJ treatment of Norway spruce appears to have practical potential as a tool for increasing plant resistance to fungal infection, but with a modest reduction in sapwood growth.     

中国红豆杉中主要紫杉烷类物质的分布研究

对中国红豆杉[Taxus chinensis(Pilg.)Rehd.]中的紫杉烷类化合物紫杉醇及两种重要前体巴卡亭III(B III)和10-脱乙酰巴卡亭III(10-DAB)的分布进行了研究.结果表明,紫杉醇的含量在同一棵树中的分布依主干皮、根皮、侧枝树皮、种子、须根、嫩枝、叶的次序递减,并发现侧枝茎中紫杉醇在横截面方向呈梯度分布,且紫杉醇含量最高部位是在次生木质层.10-DAB和B III在叶中含量最高,在须根中含量最低,叶子中10-DAB的含量达0.02%～0.03%,高于B III的含量.在2～3 a生的小树枝、叶中未检测到紫杉烷类物质,提示其合成可能与植株发育有关.紫杉醇在30 a 树龄的当年生嫩叶中春季高于秋季,而10-DAB在当年生和多年生的叶中积累都较多,且秋季比春季含量高.     

Anatomical and FTIR analyses of phloem and xylem of <Emphasis Type="Italic">Tetracentron sinense</Emphasis>

The fast growth of Tetracentron sinense is a potential valuable timber resource, but whether its anatomy and chemical components are suitable for timber is unknown. We used light microscopy and SEM to examine the anatomical structure and FITR to measure the chemical components of the phloem and xylem of this tree. Radial variations in growth ring width and tracheid dimensions were also evaluated. The sieve tube, phloem parenchyma cell and sclereids clusters were the main cells in phloem, and the tracheid was the fundamental cell in xylem. An unusual tracheid type, fiber-tracheids or vessel-liked elements was visible. Wood rays nonstoried, uniseriate and multiseriate, including heterogeneous II, occasionally I, and usually 3–6 cells wide. The mean growth-ring width was 2.53 ± 0.46 mm, and the percentage of late wood was over 60%. For radial variation, growth-ring width increased at an early growth stage, and reached the largest increment during years 11–15, then decreased. The maximum growth-ring width was 5.313 mm. During late growth (60–85 years), trees also maintained a high radial growth increment. Radial variation in the percentage of late wood was uniform, about 50–70%, throughout the growth years. Growth patterns in the length and width of early and late wood were similar as the trees aged. From the FTIR results, the chemical components differed significantly between xylem and phloem, hemicellulose in particular was higher in the xylem than in the phloem, where it was apparently absent. All of these suggest that the composition of phloem in T. sinense is very similar to that of hardwood, and it has higher growth ratio and uniform wood properties.     

Foliar mineral composition, fertilization and dieback of Norway spruce in the Belgian Ardennes

Needles from healthy Norway spruce (Picea abies (L.) Karst.) at Willerzie in the West Ardennes and from trees with symptoms of dieback at Langesthal in the East Ardennes were analyzed by age class for mineral composition. Both stands were on acid oligotrophic soils. At Willerzie, needles were sampled from plots fertilized 12 to 17 years earlier (dolomitic lime plus N, P and K) as well as unfertilized plots. Effects of fertilization included increased levels of calcium, manganese, phosphorus, and copper and reduced levels of total sulfur, sulfate-S, sulfate-S:total S, potassium and aluminum. Levels of calcium, magnesium, copper and boron were low at both sites and, at Langesthal, calcium and magnesium may have been deficient. Sulfur level was normal at Willerzie, but at Langesthal, mean sulfur content for needles of all age classes was 198 mg 100 g(-1) dry weight, a level that may be toxic. In older needles, the N:S ratio at Langesthal was below the threshold value of eight reported to be necessary for healthy growth. Other symptoms of stress observed were high sulfate-S:total S and nitrate-N:total N ratios. At Langesthal, manganese level was probably adequate although only one-fifth the level at Willerzie. Levels of aluminum and iron were very high at both sites. Most of the iron and much of the aluminum occurred as a surface deposit that could be removed by washing the needles in chloroform.     

Freezing pattern and frost killing temperature of apple (Malus domestica) wood under controlled conditions and in nature

The freezing pattern and frost killing temperatures of apple (Malus domestica Borkh.) xylem were determined by differential thermal analysis and infrared differential thermal analysis (IDTA). Results from detached or attached twigs in controlled freezing experiments and during natural field freezing of trees were compared. Non-lethal freezing of apoplastic water in apple xylem as monitored during natural winter frosts in the field occurred at -1.9?±?0.4 °C and did not change seasonally. The pattern of whole tree freezing was variable and specific to the environmental conditions. On detached twigs high-temperature freezing exotherms (HTEs) occurred 2.8 K below the temperature observed under natural frosts in the field with a seasonal mean of -4.7?±?0.5 °C. Microporous apple xylem showed freezing without a specific pattern within a few seconds in IDTA images during HTEs, which is in contrast to macroporous xylem where a 2D freezing pattern mirrors anatomical structures. The pith tissue always remained unfrozen. Increasing twig length increased ice nucleation temperature; for increased twig diameter the effect was not significant. In attached twigs frozen in field portable freezing chambers, HTEs were recorded at a similar mean temperature (-4.6?±?1.0 °C) to those for detached twigs. Upon lethal intracellular freezing of apple xylem parenchyma cells (XPCs) low-temperature freezing exotherms (LTEs) can be recorded. Low-temperature freezing exotherms determined on detached twigs varied significantly between a winter minimum of -36.9 °C and a summer maximum -12.7 °C. Within the temperature range wherein LTEs were recorded by IDTA in summer (-12.7?±?0.5 to -20.3?±?1.1 °C) various tiny clearly separated discontinuous freezing events could be detected similar to that in other species with contrasting XPC anatomy. These freezing events appeared to be initially located in the primary and only later in the secondary xylem. During the LTE no freezing events in the bark and central pith tissue were recorded. Attached twigs were exposed to various freezing temperatures at which LTEs occur. Even if 60% of XPCs were frost-damaged twigs were able to recuperate and showed full re-growth indicating a high regeneration capacity even after severe frost damage to XPCs.     

Root and transpiration studies on young Norway spruce trees with die back symptoms in Sweden

In the spring of 1977, scattered trees in young plantations of Norway spruce showed discoloration and loss of 1976 needles on twigs just below the apex. The main roots of damaged trees were smaller than those of healthy trees, and the extension roots lacked short roots and mycorrhizae. The transpiration rate of the current shoots of damaged trees was also higher. The damage was assumed to be the result of drought during the two previous years; the trees were thought to differ genetically in their ability to withstand water stress.     

Can the structure of dormant cambium and the widths of phloem and xylem increments be used as indicators for tree vitality?

We investigated the structure and width of the dormant cambium and of the increments of phloem and xylem of Quercus robur to estimate their potential as indicators for tree vitality. The samples were taken from three woodlands, two in Slovenia [Krakovo forest (KRA) and Murska Suma (MUS)] and one in Croatia [Kobiljak (KOB)], with reported tree decline. The number of dormant cells seems to reflect the initial capacity of the cambium to accomplish cell division. With the exception of two trees at KRA, cell production was always higher on the xylem side than on the phloem side. The annual phloem increments were narrower, less variable among trees and with clear lower and upper limits. With increased cambial cell productivity, the share of the xylem in the total annual radial increment increased following a curvilinear function. In trees with an annual radial increment >3.5 mm, the xylem size represented more than 90 % of the total radial growth. The anatomical variables analyzed show that the most limiting environmental conditions seem to prevail at KRA, whereas the conditions at MUS seem to be most favorable in terms of radial growth. Analysis of the width and structure of xylem and phloem increments, the number of dormant cambial cells and their inter-relationships can provide additional information on the vitality of oaks.     

Photosynthetic traits and antioxidative defense responses of Pinus yunnanensis after joint attack by bark beetles Tomicus yunnanensis and T.minor

Bark beetles Tomicus yunnanensis and T. minor are two important pests of Pinus yunnanensis and can cause massive death of pine trees. In this study, we examined several traits related to photosynthesis in P.yunnanensis and their relationship with antibiotic defense responses after joint attack by the two bark beetles at the shoot and the trunk stages. When shoots were attacked by the beetles, the abundance of chlorophylls, carotenoids, and the rates of net photosynthesis(Pn) and transpiration(E) decreased in needles, while the levels of superoxide dismutase and malondialdehyde remained unchanged in both needles and phloem. The activity of peroxidases also remained unchanged in needles, but increased in phloem.The activity of catalases increased in both needles and phloem. When trunks were attacked by the bark beetles,chlorophyll abundance, Pn, E, and antioxidative enzyme activities all declined, and the declines were more pronounced than in the attacked shoots. A decrease in protein concentrations was also observed in needles and phloem from the attacked pines. Attack on shoots by the bark beetles suppressed host defense and provided a favorable environment for larval growth and development, resulting in long-term decline of pine growth potential. The results suggest that attacks on trunks by beetles caused more severe damage to host trees than attacks on shoots.     

Acid fog removes calcium and boron from fir tree: One of the possible causes of forest decline

We have observed acid fog in Mt. Oyama adjacent to the Kanto Plains in Japan, where acid fog under pH 3 has been frequently observed and natural fir forest (Abies firma) has been declining. We applied the simulated acid fog to the needles of fir twigs and the treatment removed calcium, boron, and cell-wall acidic sugars from the needles. We also observed high contents of calcium, boron, and acidic sugars in throughfall collected under the declining forest canopy. Ca⁺⁺ and Mg⁺⁺ ion leach via ion-exchange with major cations of fog-water and the amount of leached boron increased with the increase in the concentration of leached Ca⁺⁺. The deficiency of calcium and boron by leaching could be one of the main causes of decline of the fir forest.     

Effects of foliar potassium concentration on morphology, ultrastructure and polyamine concentrations of Scots pine needles

We examined the effects of three foliar potassium concentrations (high, intermediate and low) on the morphology, ultrastructure and polyamine concentrations of current-year and 1- and 2-year-old needles of 30-year-old Scots pine (Pinus sylvestris L.) trees. Foliar K concentration had only a slight effect on needle morphology. The sclerenchyma cell walls were thinner, the xylem area was larger, and the resin ducts were smaller in needles with a low K concentration than in needles with a high or intermediate K concentration. In addition, the bundle sheath cells were collapsed in needles having a low K concentration. The secondary growth of phloem tissue and the mesophyll area were greater in needles with a high or intermediate K concentration than in needles with a low K concentration, possibly indicating greater production of photoassimilates in these trees. At the ultrastructural level, mesophyll cells with enlarged central vacuoles and small vacuoles containing electron-dense material were common in needles having a low K concentration. Enlargement of the central vacuole coincided with an exponential increase in putrescine concentration in needles with a low K concentration, suggesting that the central vacuole may function as a storage site for putrescine.     

Differences in composition of endophytic mycobiota in twigs and leaves of healthy and declining Quercus species in Italy

The composition of the endophytic fungal assemblages in Quercus cerris, Q. pubescens and Q. robur, three oak species most susceptible to decline, was investigated in Fagarè (Padova), Ulignano (Pisa), and Radda in Chianti (Firenze). The endophytic assemblages were studied as a function of health status of the whole tree and of twigs and leaves. The effect of station altitude was tested on Q. cerris. Samples were gathered from healthy and declining trees. In April (bud opening time) 20 twigs and 10 leaves were collected from each tree. Fungi were isolated from twig segments and leaf portions. A total of 23 fungal genera with 27 different species were isolated, in addition to a few sterile mycelia. Eleven species were found to be common to all three oak species, while other species were found only on one oak species, suggesting host specificity. Among the fungal species isolated, some were known to be pathogenic (Apiognomonia quercina, Colpoma quercinum, Diplodia mutila, Phomopsis quercina). The colonization frequency (CF) of pathogenic species varied between 0.9% for A. quercina in Q. cerris and 60.2% for P. quercina in Q. robur. Analysis of variance showed a statistically significant difference among the oak species tested. The CF was higher in declining trees in comparison with healthy trees, and also in twigs vs. leaves. The CF was found to be higher for Q. cerris trees growing at sea level as compared with those situated at an elevation of 350–400 m. In stands where, on account of particular ecological conditions, pathogenic behaviour is displayed simultaneously by more than one endophytic fungal species, trees accentuate their decline.     

Do increasing respiratory costs explain the decline with age of forest growth rate?

Once forests have achieved a full canopy, their growth rate declines progressively with age. This work used a global data set with estimates from a wide range of forest types, aged 20-795 years, of their annual photosynthetic production(gross primary production, GPP) and subsequent above-plus below-ground biomass production(net primary production, NPP). Both GPP and NPP increased with increasing mean annual temperature and precipitation. GPP was then unrelated to forest age whilst NPP declined progressively with increasing age. These results implied that autotrophic respiration increases with age. It has been proposed that GPP should decline in response to increasing water stress in leaves as water is raised to greater heights as trees grow taller with age. However, trees may make substantial plastic adjustment in mor phology and anatomy of newly developing leaves, xylem and fi ne roots to compensate for this stress and maintain GPP with age. This work reviews the possibilities that NPP declines with age as respiratory costsincrease progressively in, any or all of, the construction and maintenance of more complex tissues, the maintenance of increasing amounts of live tissue within the sapwood of stems and coarse roots, the conversion of sapwood to hear twood, the increasing distance of phloem transport, increased turnover rates of fine roots, cost of supporting very tall trees that are unable to compensate fully for increased water stress in their canopies or maintaining alive competitively unsuccessful small trees.     

Organic and inorganic sulfur transport in the xylem sap and the sulfur budget of Picea abies trees

Temporal changes in inorganic and organic sulfur compounds (sulfate, glutathione, cysteine, methionine) were analyzed in xylem sap of 40-year-old Norway spruce (Picea abies (L.) Karst.) trees growing on acidic soils at a healthy and a declining stand in the Fichtelgebirge (North Bavaria, Germany). Studies were carried out (1) to quantify glutathione (GSH) transport in the xylem of spruce, (2) to study the significance of reduced sulfur versus sulfate (SO(4) (2-)) transport in the xylem, and (3) to compare total sulfur (S) transport in the xylem with the amount of foliar uptake of SO(2) in an air-polluted environment. Glutathione was the main reduced S compound in the xylem ranging in concentration from 0.5 to 5 &mgr;mol l(-1). Concentrations of inorganic SO(4) (2-) in the xylem sap were up to 50 times higher than those of GSH ranging from 60 to 230 &mgr;mol l(-1). During the growing season, concentrations of all S compounds in the xylem were highest in May (up to 246 &mgr;mol l(-1)) and decreased during summer and fall (up to 21 &mgr;mol l(-1)). On average, SO(4) (2-) concentrations in xylem sap were 30% higher at the declining site compared with the healthy site. Diurnal changes in organic S compounds were significant for GSH and cysteine with high concentrations during the night and low concentrations during the day. Diurnal changes in inorganic concentrations were not significant. Xylem sap concentrations of SO(4) (2-) and cysteine were twice as high and GSH concentrations were tenfold higher in surface roots than in branches. At both sites, transport of organic S was low (up to 3% of total S) compared to transport of SO(4) (2-). Annual transport of total S in the xylem (SO(4) (2-) was the main component) ranged from 60 to 197 mmol tree(-1) year(-1) at the healthy site and from 123 to 239 mmol tree(-1) year(-1) at the declining site. Although gaseous uptake of SO(2) was estimated to be similar at both sites (38 mmol tree(-1) year(-1); Horn et al. 1989), the ratio between annual gaseous uptake of SO(2) and transport of S in the xylem was 1:4 and 1:5 at the healthy and declining sites, respectively.     

日本松干蚧口针及其危害枝条切片观察

日本松干蚧(Matsucoccus matsumurae Kuwana)发生数十年以来,各地均进行了研究和防治,取得了成果和效益。而研究日本松干蚧口针和吸食部位以及损害松树的原因尚未见有报导。笔者自1985年起开展这一研究,现将初步结果整理如下。