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1.
Light availability and infestation by the green spruce aphid (Elatobium abietinum) are key factors affecting the growth of Sitka spruce (Picea sitchensis) seedlings under a mature tree canopy, but their combined effect on seedling growth has not previously been quantified. A controlled outdoor experiment in which light levels (high light (HL): 100%, intermediate light (IL): 24%) and aphid infestation (absence/presence) were manipulated was conducted over 2 years to look at the effects on seedling growth and biomass distribution patterns. Aphid population assessments showed a significantly increased population density under IL, with three to four times higher cumulative aphid densities than that under HL. Defoliation rates of infested seedlings were directly related to aphid density. Total seedling biomass was strongly reduced in IL, and aphid infestation caused additional reductions in the biomass of particular components of the seedlings. Dry weight (DW) of older (≥1-year-old) needles in infested trees was significantly decreased in both years. Total root DW at the end of the second year was significantly affected by aphid infestation, and the reduction (14-18%) was similar in IL and HL treatments despite large differences in aphid density. Biomass distribution patterns in infested trees were similar to that of uninfested trees within each light treatment, indicating that the relative decreases in root biomass were accompanied by similar reductions in distribution to the above-ground parts of the seedlings. Leader extension growth of infested seedlings was reduced by 15-17% compared with uninfested seedlings under IL, whereas only a 2-3% reduction in leader extension of infested seedlings under HL was observed. The results showed that the response of seedlings to E. abietinum were primarily dependent on the light environment. The significant reduction caused by aphids on the total DW of older needles and roots, and on leader extension growth, does suggest the potential for effects to accumulate over time.  相似文献   

2.
The needles of pine trees are indicative of the overall health of the tree, and their length is affected by many factors. This study describes the effect of high levels of pathogen infection on lengths of both needles and shoots. Dothistroma septosporum is an important foliar pathogen of pines causing necrosis and premature defoliation with successive years of high infection leading to growth reduction and in extreme cases tree death. Corsican pine trees with all foliage infected by D. septosporum had needles of primary, secondary and tertiary shoots reduced by 31.8%, 44.0% and 64.2%, respectively, compared to non‐infected trees. Needle lengths were reduced both in the upper and lower portions of the canopy, with a greater reduction lower in the canopy. Trees with high levels of infection had shorter shoots in the upper canopy with fewer, shorter needles on these shoots compared to trees with low levels of infection. The results demonstrate the substantial negative effect on needle and shoot lengths of trees with high levels of D. septosporum infection, comparable to factors such as water and nutrient availability known to have a strong influence on these parameters. The reductions in length reduce the photosynthetic capacity of the tree and compound the immediate reductions caused by necrosis and premature defoliation. These effects on needle and shoot lengths contribute to the reductions in volume growth of affected trees and, moreover, are longer lasting than the immediate effects of necrosis and premature defoliation.  相似文献   

3.
This study represents a geographical approach to morphological variation of cones, needles and seeds in natural dragon spruce (Picea asperata Mast.) populations across western China and the relationship between morphological traits and weather variation for the origins of populations. Significant differences were found for all cone, needle and seed traits among populations and trees within populations. The coefficients of variation in 2 higher latitude populations were higher than those in other populations in all traits except for needle width and seed width. Fairly high within-population variability was also detected for cone dry weight, seed dry weight and needle traits in this study. Cone length, cone width, cone dry weight, seed scale length, seed scale width and seed wing length were strongly positively intercorrelated. Seed length and seed dry weight showed a significantly positive correlation with seed wing length. And needle width showed a significant negative correlation with cone traits, seed scale length, seed wing length and seed length. The higher the annual mean temperature, the higher the average cone length. This study also revealed a negative linear dependence on latitude: the higher the latitude, the lower the population average seed length. Seed dry weight was positively and linearly related with altitude.  相似文献   

4.
Needle development and shoot growth were studied in 14- and 20-year-old Sitka spruce (Picea sitchensis (Bong.) Carr.) planted on an oligotrophic peat and fertilized with N, P and K (control), N and P (-K), N and K (-P) or unfertilized (-NPK). Shoot extension, needle number, and the size and number of needle cells were observed throughout the season. Beginning with bud burst in early May, needle growth continued for nine weeks in the control treatment. Needle growth in the mineral deficiency treatments terminated at the same time as in the controls, although it did not begin until up to three weeks later than in the control trees. Needles developed acropetally along the shoot, with basal needles completing their development first. Cell division in needles of control trees lasted about 3 weeks, with cell expansion continuing for a further 3 weeks. Initial mean cell cycle times were shortest for proximal needles following bud burst, but lengthened as the season progressed. Cell number increased from 6600 in the primordia to 200 000 in mature needles. Final needle dimensions and cell number varied according to the position of the needle on the shoot. The largest needles with the most cells were near the middle of the shoot. Relative to the controls, all mineral deficiency treatments reduced shoot length, bud and needle dimensions, and needle cell number, particularly the -P treatment. Potassium and P deficiency treatments reduced primordial cell number by up to 42%, whereas final mean cell size was 30% greater in the -P treatment, 17% greater in the -NPK treatment, but 14% smaller in the -K treatment. Mean cell size was constant for needles at all positions in any particular treatment at any time, so that final needle size was determined by cell number alone. Needle dry weight/fresh weight ratio continued to increase until early August, with significantly higher ratios in the -K and -P treatments than in the controls.  相似文献   

5.
Forest decline has been attributed to the interaction of several stressors including biotic factors such as mistletoes and climate-induced drought stress. However, few data exist on how mistletoes are spatially arranged within trees and how this spatial pattern is related to changes in radial growth, responses to drought stress and carbon use. We used dendrochronology to quantify how mistletoe (Viscum album L.) infestation and drought stress affected long-term growth patterns in Pinus sylvestris L. at different heights. Basal area increment (BAI) trends and comparisons between trees of three different infestation degrees (without mistletoe, ID1; moderately infested trees, ID2; and severely infested trees, ID3) were performed using linear mixed-effects models. To identify the main climatic drivers of tree growth tree-ring widths were converted into indexed chronologies and related to climate data using correlation functions. We performed spatial analyses of the 3D distribution of mistletoe individuals and their ages within the crowns of three severely infested pines to describe their patterns. Lastly, we quantified carbohydrate and nitrogen concentrations in needles and sapwood of branches from severely infested trees and from trees without mistletoe. Mistletoe individuals formed strongly clustered groups of similar age within tree crowns and their age increased towards the crown apex. Mistletoe infestation negatively impacted growth but this effect was stronger near the tree apex than in the rest of sampled heights, causing an average loss of 64% in BAI (loss of BAI was ～51% at 1.3 m or near the tree base). We found that BAI of severely infested trees and moderately or non-infested trees diverged since 2001 and such divergence was magnified by drought. Infested trees had lower concentrations of soluble sugars in their needles than non-infested ones. We conclude that mistletoe infestation causes growth decline and increases the sensitivity of trees to drought stress.  相似文献   

6.
We investigated current shoot properties in two contrasting vertical positions (leader crown; LC, and lower branch; LB) within the crowns of mature trees of two subalpine conifer species, Abies mariesii and A. veitchii. For both LCs and LBs, shoot length decreased with increasing branching order. However, shoot properties were different between LCs and LBs. Shoots in LCs had more needle biomass per unit of shoot length. Shoots in sunny conditions pack needles closer along the shoot and intercept incoming light more completely. This causes the shoots in the LCs to have more needles. In contrast, less needle packing per unit shoot length in LBs results in the avoidance of mutual shading among needles in order to intercept limited light more effectively. Because branch systems in lower layers tend to be more shaded, the quantity of irradiance received by the shoots in LBs is smaller. Thus, reduced needle amounts on the shoots in LBs reflect the needle arrangement acclimating to the lower light availability. This study suggests the importance of changes in the properties of individual shoot as a component of a branch system and accordingly a whole-crown system in mature canopy trees of A. mariesii and A. veitchii.  相似文献   

7.
Green spruce aphid (Elatobium abietinum) feeds primarily on the 1-year-old and older needle leaves of spruce (Picea spp.) and is a major defoliator of commercial Sitka spruce plantations in the UK and other maritime regions of north-west Europe. The impact of E. abietinum on mid-rotation (23-28-year-old) Sitka spruce in Radnor Forest, in mid-Wales, was determined by comparing aphid population densities, defoliation and growth between plots of trees that were either treated with insecticide or were left untreated. The experimental treatments were maintained for 5 years and over this period (2002-2006) peak E. abietinum densities in the untreated plots varied from 5 to 36 aphids per 100 needles. These densities, which were representative of low to moderate rates of infestation, were associated with low rates of defoliation (0-8%), but they had a significant (P ? 0.01) impact on mean radial increment (RI) and mean volume increment (VI). In 2005, the year with the highest aphid populations, peak densities in the untreated plots averaged 14 aphids per 100 needles and this rate of infestation reduced RI by 17%, VI by 10% and the dry weight of current-year needles by 10%. On average, across all years, infestation by E. abietinum reduced mean annual VI by 6%. Comparisons with previous studies on the impact of the aphid on 4-year-old, 7-year-old and 9-year-old Sitka spruce (Straw et al., 2005) indicate that defoliation by E. abietinum, on a per capita basis, decreases as trees grow older, but that the impact on VI increases. The different growth response of young and mid-rotation Sitka spruce to infestation is related to differences in canopy structure, particularly in the ratio of current-year needles to older needles, and the greater demands on photosynthetic production in older trees that arise from the need to support an increasing quantity of non-photosynthetic structural tissues in branches, stem and roots.  相似文献   

8.
Investigations on I. typographus populations were conducted in 1992–1994 in the Sudeten (south-western Poland), in Norway spruce stands that were devastated previously by insect outbreaks and that exhibited severe forest decline. Crown defoliation and the volume of infested trees per hectare were used as indices of stand health and breeding conditions; variability in the infestation density, gallery length, and body length of beetles were parameters that were used to estimate the response of I. typographus populations to changing stand conditions. In stands located in the zone above 800 m a.s.l. where both the level defoliation and tree mortality were higher, the infestation density and body length of I. typographus were higher than in the zone below 800 m a.s.l., where stands were less damaged. Contrarily, the gallery length was higher in lower montane zone, where lower infestation densities and body length values were found. A proposed mechanism for this interaction, which is attributed mainly to better breeding conditions for I. typographus populations in weakened stands, is presented and discussed.  相似文献   

9.
Vitellaria paradoxa, known as Shea tree or karité, is one of the dominant trees of agroforestry parklands in Sahelian region with an immense commercial value. In this study, we examined infestation of fruit-bearing shoots and fruits on two sites in western Burkina Faso across different reproductive phenophases, crown heights and aspects. The number of infested and healthy shoots and fruits were counted on 60 reproductively active trees randomly selected from on-year shea trees in each site, using a metal frame (0.5 × 0.5 m) to mark a fixed area of the canopy for sampling. The shoot and fruit borer was identified as Salebria sp. (Lepidoptera:Pyralidae), which was not reported earlier. The proportion of infested trees and shoots varied significantly between sites and among reproductive phenophases (P < 0.0001). The site close to the natural forest had the highest infestation rate than distantly located site. Among reproductive phenophases, the proportions of infested trees and shoots were the highest during flowering and fruit set, respectively. Fruit infestation also varied significantly between sites, crown heights and aspects (P < 0.001). The site close to the natural forest and the crown with north-east aspect were heavily infested while the bottom part of the crown had the lowest infestation. As a whole, 49–80% of trees, 24–31% of shoots and 4–15% of fruits were found infested, depending on the site. These infestation levels combined with precocious flower and fruit abscissions would pose serious economic losses that justify a call for immediate pest control initiatives.  相似文献   

10.
Concentrations of amino acids, total amino nitrogen, and phenols, ratio of phenol to amino nitrogen, and pH in the bark of American beech, Fagus grandifolia, were determined to see if these chemical constituents were correlated with susceptibility of trees to infestation by the beech scale, Cryptococcus fagisuga. The relationship of these bark constituents with scale infestation levels (none, light, and moderate), tree size (large vs. small trees), and bark shading was determined. Levels of some individual amino acids and total amino acid content in moderately infested trees were significantly higher than in uninfested trees. Large uninfested beech trees, which tend to be more readily infested by scale, had significantly higher concentrations of aspartic acid. Concentrations of phenol in the outer bark of both infested and uninfested trees were significantly higher than in the inner bark. However in bark infected by Nectria, phenol levels in the inner bark were significantly higher than in the outer bark.  相似文献   

11.
Arceuthobium sichuanense is a hemiparasitic angiosperm that infects Qinghai spruce (Picea crassifolia Kom.) in Qinghai province, China, and causes severe damage to spruce forests in Qinghai‐Tibet Plateau. In this study, the impact of A. sichuanense infection on mature and young trees of Qinghai spruce was evaluated by examining needle and current‐year shoot morphology, needle water and nitrogen‐use efficiency (NUE) and needle nitrogen concentration. The most apparent effect of A. sichuanense infection was a significant reduction in both needle size distal to infection and current‐year shoot length in the infected branches (p < 0.001). Per cent reductions in needle and current‐year shoot length were similar between mature and young trees (58.9 vs. 56.3%; 59.7 vs. 62.9%). There was a high degree of correlation in foliar δ15N values between the dwarf mistletoe and its host trees (R2 = 0.9017, p < 0.001), while the foliar δ13C values of A. sichuanense were similar to those of infected mature and young spruce trees. The dwarf mistletoe infection also resulted in a significant decrease in host needle N concentration and δ13C values (p < 0.001). The per cent reduction in needle N concentration in young trees was nearly twice as much as that in mature trees (20.49 vs. 11.54%), while the per cent reduction in needle δ13C values was similar between young and mature trees (?0.98 vs.?1.1‰). The NUE in mature trees was not affected by A. sichuanense infection, but the NUE in young trees was increased by the infection.  相似文献   

12.
Studies were carried out in 1999, 2005 and 2007 in the area of Kunda cement plant in Northeast Estonia on sample plots 3 km W and 2.5 and 5 km E of Kunda. As control stands, two plots for pine and spruce were established in Lahemaa National Park (34-38 km W of Kunda). The selected pine and spruce stands were 75-85-year-old Myrtillus site type, of 0.7-0.8 density and II quality class, with moderately dense or sparse understorey. The values concerning needle density and number of needle scars were higher for shoots formed in the period of higher pollution than for the shoots grown under a considerably lower pollution load. Although the cement dust pollution has notably decreased from year to year, the number of needle pairs per 1 cm of the shoot was 1.8-2.1 times greater in the shoots formed in 1998 than in those formed in 2003, whereas the changes were statistically reliable. Possibly the low temperatures at the time of shoot and needle formation affected the density of needles on all sample plots, and thus the number of needles on shoots formed in 2003 was many times smaller. After the significant fall in the pollution load since 1996 the length growth of needles intensified around Kunda cement plant, at the same time no changes occurred in the length growth of needles in the control area. As compared to the data from 1998, the length growth of pine needles had improved, especially 2.5 and 5 km E from the cement plant, needles being respectively 1.5 and 1.1 cm longer than 6 years ago. The stimulation of the growth of pine and spruce needles 2.5 and 5 km E of the cement plant may be a sign of a positive effect of reduced doses of cement dust in soil. The greater length of pine and spruce needles is the reason for the larger biomass of the needles.  相似文献   

13.
This study represents a geographical approach to morphological variation of cones, needles and seeds in natural dragon spruce （Picea asperata Mast.） populations across western China and the relationship between morphological traits and weather variation for the origins of populations. Significant differences were found for all cone, needle and seed traits among populations and trees within populations. The coefficients of variation in 2 higher latitude populations were higher than those in other populations in all traits except for needle width and seed width. Fairly high within-population variability was also detected for cone dry weight, seed dry weight and needle traits in this study. Cone length, cone width, cone dry weight, seed scale length, seed scale width and seed wing length were strongly positively intercorrelated. Seed length and seed dry weight showed a significantly positive correlation with seed wing length. And needle width showed a significant negative correlation with cone traits, seed scale length, seed wing length and seed length. The higher the annual mean temperature, the higher the average cone length. This study also revealed a negative linear dependence on latitude： the higher the latitude, the lower the population average seed length. Seed dry weight was positively and linearly related with altitude.  相似文献   

14.
In this study the needle nutrient status of Sirococcus shoot blight‐diseased and healthy Norway spruces (Picea abies) was investigated. Element contents were determined in the current year and 3‐year‐old needles collected in December 2000 from 72 mature trees randomly distributed in the investigated stand. Half of the trees were severely affected by Sirococcus conigenus and had a reduced crown whereas the other trees were unaffected and vigorous. Compared with the uninfected trees, contents of Mg, Ca, P and Mn were significantly lower in the current year and 3‐year‐old needles of diseased trees. Furthermore, analyses of the nutrient contents of needles from severely affected trees revealed insufficient Mg and Ca supply and enhanced N/Mg and N/Ca ratios, when compared with threshold values. The results of this study suggest that imbalanced tree nutrition either has a significant impact on the expression of Sirococcus disease symptoms or fungal infection itself is affecting needle nutrient contents.  相似文献   

15.
SEABY  D. A.; MOWAT  D. J. 《Forestry》1993,66(4):371-379
Following a severe attack by Elatobium abietinum (Walker) in1979–80, annual observations were made for 5 years on320 pole-size Sitka spruce in a nitrogen and phosphate experimentlaid out on oligotrophic blanket bog. Leader length was reduced by 25.5 per cent in 1980–81compared with the 3 previous years. This reduction was highlycorrelated with defoliation, being 39.4 per cent for trees thatlost all 1977–79 needles and 8.4 per cent in trees thatlost no needles. There were strong interactions with fertilizertreatment. Defoliated trees continued to grow less well duringthe 5 years of observations and were more likely to go intogrowth check. Early flushing trees suffered significantly less growth reductionfollowing defoliation. The same trees consistently broke budearly over 4 years of observations. Early bud burst was notcorrelated with height, girth or defoliation. No reattack by aphids was observed in the experiment until 1985–86and in this minor infestation the trees previously colonizedwere found to be significantly more likely to be recolonized. A novel measuring-rod for tree height is described.  相似文献   

16.
Samples of Norway spruce (Picea abies [L.] Karst) were dendrochronologically investigated in order to detect infestations by Dendroctonus micans (Kug.), the great spruce bark beetle (Col. Scolytidae), a relatively recent introduction to France. Uninfested natural forests located in the north-eastern French Alps and heavily infested plantations in the Ardèche region (Massif Central) were compared. The penetration holes bored in trunks by the bark beetle induced visible marks on wood, such as extreme ring width reductions, locally missing rings and crescent-shaped resin patches between consecutive rings that make possible a post-infestation dating.The outbreak began in 1979, 5 years prior to first insect visual detection by foresters. In the infested forest, tree basal area growth was not as sustained as in uninfested natural stands, but showed an inflection point at an unusually young tree age (from 30 to 40 years). Ring widths showing extreme synchronous radial growth reductions were caused either by excessively cold periods (e.g. in 1948, 1980, 1984, 1992) or by summer drought (as in 1986). Most of these weak growth years were shared with uninfested sites. In healthy forests, the consequences of extremely cold years were usually recorded only in high elevation stands, especially near the timberline, whereas summer drought effects were mostly visible in low altitude forests. By contrast, both phenomena were recorded in the infected Ardèche plantation. An analysis of tree-rings and monthly climate confirmed that Norway spruce growth in Ardèche plantations was reduced by excessively low minimum temperature during most parts of the year prior to ring formation, by higher than average maximum temperature during current spring and summer, and by drought in winter, spring and summer. Thus, the regional Ardèche climate with both cold winters and dry summers (especially in July) seems to weaken spruce trees planted there. Moreover, tree sensitivity to climate was found to be greatly enhanced by insect infestation. Such interactions between climatic stress and insect outbreak led to forest dieback in a 15–20-year period, when trees were still young (less than 70 years), and without any tree recovery. Therefore, in that region spruce plantations should be replaced by non-host species of Dendroctonus micans, especially where soil conditions may exacerbate drought effects.  相似文献   

17.
Relationships between tree mortality and bark beetle infestation onAbies veitchii at the wave-regenerated forest in Mt. Asahi, Okuchichibu area were investigated. Most of the firs with green needles and newly developed current year’s shoots in the dieback zone were heavily infested by bark beetles before the death of the trees. After heavy infestation of beetles, about half of the infested firs died within the year, and the other half died in the next year. When the species composition of bark beetles and associated ophiostomatoid fungi were investigated in Mt. Asahi and also at a typical wave-regenerated forest in Mt. Shimagare, Yatsugatake area,Cryphalus montanus andC. piceae were dominant beetle species for Mt. Asahi and Mt. Shimagare, respectively.Ophiostoma subalpinum andO. europhioides were dominant fungal species at both wave-generated forests. BecauseO. subalpinum was detected more frequently from deeper areas of sapwood thanO. europhioides, it was suggested that the fungal species may accelerate the death of stressed firs in wave-regenerated forests. Contribution No. 169, Laboratory of Plant Parasitic Mycology, Institute of Agriculture and Forestry, University of Tsukuba.  相似文献   

18.
Both drought and root pruning (RP) increased the number of cones induced when black spruce (Picea mariana (Mill.) B.S.P.) grafts were injected with gibberellins A(4/7) (GA), but their effects on predawn shoot water potential and current-year needle development differed. Drought decreased predawn shoot water potential (Psi(pd)), but only during the period when irrigation was withheld, and it had no effect on the growth or gas exchange properties of current-year needles. Conversely, root pruning had little effect on Psi(pd), but it resulted in trees with smaller current-year needles that had lower nitrogen and chlorophyll concentrations and reduced rates of gas exchange up to the later stages of shoot elongation compared with needles of control trees. These findings are discussed in relation to potential effects on the development of induced cones in the following growth cycle.  相似文献   

19.
To trace any possible effects of air pollution stress and injuries on trees around a city centre, needles of Scots pine (Pinus Syivestris L.) and Norway spruce (Picea abies (L.) Karst.) from sites of different distances and directions from Stockholm were analysed. Comparisons were made between needles of different ages, a half to three and a half years old (C ‐ C + 3) for pine up to six and a half years old (C ‐ C + 6) for spruce. Epicuticular waxes, studied by scanning electron microscopy (SEM), showed age‐dependent but site‐independent changes. Different age‐patterns were found in the pine and spruce needles but no dust coverings or lesions were detected, irrespective of the sites. A proportionally increasing peroxidase activity (EC 1.11.1.7) with increasing needle‐age up to the third year was observed in both species. No certain indication of site‐effects, expressed by an unproportional increase of the enzyme activity with increasing needle‐age could be found. Chlorophyll a and b, phaeophytin a and b and carotenoid analyses on the corresponding materials only indicated disturbances in trees from some scattered sampling plots. Thus, neither could the pigment analyses reveal any signs of a general pollution stress in pine and spruce trees from the area in question.  相似文献   

20.
We compared shoot and needle morphology and photosynthesis in Picea glehnii (Friedr. Schmidt) M.T. Mast. and Picea jezoensis (Sieb. et Zucc.) Carr. trees planted on immature volcanic ash and well-developed brown forest soils to investigate whether species differences in morphological and physiological plasticity affected tree growth on different soil types. Height growth of P. glehnii was reduced by about 10% on volcanic ash compared with forest soil, whereas that of P. jezoensis was reduced by more than 60%. Needle morphology of P. glehnii was unaffected by soil type. In contrast, needles of P. jezoensis trees growing on volcanic ash were shorter, narrower and thicker, with less dry mass and area, than those of trees growing on forest soil, and specific needle area was lower, indicating lower foliar light-interception efficiency. In both species, changes in needle morphology with increasing irradiance were similar in trees growing on both soil types, indicating that plasticity of needle morphology was unaffected by soil type. In both species, shoot mass and shoot silhouette area were lower and needle mass per unit shoot mass was higher in trees growing on volcanic ash than in trees growing on forest soil. Trees of both species had more needles per unit shoot length, lower shoot silhouette to projected needle area ratios and lower shoot silhouette areas per unit shoot mass (SAM) on volcanic ash than on forest soil, indicating lower shoot-level light-interception efficiency. For P. glehnii, the response of shoot morphology to increasing irradiance was similar on both soil types, with the exception of SAM, which showed lower plasticity in trees growing on volcanic ash. In contrast, shoot-level morphological plasticity of P. jezoensis was reduced in trees growing on volcanic ash. Light-saturated maximum photosynthetic rate (P(max)) of P. glehnii was unaffected by soil type, whereas mass-based P(max) of P. jezoensis was lower in trees growing on volcanic ash than in trees growing on forest soil. In P. jezoensis trees growing on forest soil, area-based P(max) increased with increasing irradiance, but this response was not observed in trees growing on volcanic ash. As a result, area-based P(max) at the top of the canopy was 39 to 54% lower in trees growing on volcanic ash than in trees growing on forest soil. Our results indicate that constraints on morphological acclimation to high irradiances may contribute to reduced height growth of P. jezoensis on volcanic ash.  相似文献   

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