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1.
We investigated mature dwarf Abies mariesii trees growing in conifer thicket–meadow parklands on a snowy subalpine plateau, where these dwarf trees are buried in the accumulated snow in winter. We focused on structural variation in the needles, shoots, and branchlets within different crown positions (leader crown vs lower crown) of the dwarf trees. In the leader crown, which appears above the snow surface earlier than the lower crown, current-year shoots and branchlets had greater total biomass, and foliage was more closely packed along the stem axis than in the lower crown, whereas current-year shoots in the leader crown had a lower needle mass ratio than in the lower crown. These results suggest that current-year shoots and branchlets in the leader crown have a specific structure that allows them to harvest more light, although construction and maintenance costs would be higher. In contrast, the structural characteristics of current-year shoots and branchlets in the lower crown efficiently concentrate incoming light by avoiding mutual shading within foliage, thus leading to increased biomass of photosynthetic needles within shoot and branchlet biomass. Such within-crown variability at various hierarchical levels from needles to branches in mature, but very dwarf, A. mariesii trees maintains the crown and allows survival within conifer clumps in areas of subalpine parklands that receive heavy snowfall.  相似文献   

2.
We studied the effects of variation in shoot structure and needle morphology on the distributions of light and nitrogen within a Pacific silver fir (Abies amabilis (Dougl.) Forbes) canopy. Specifically, we investigated the role of morphological shade acclimation in the determination of resource use efficiency, which is claimed to be optimal when the distribution of nitrogen within the canopy is directly proportional to the distribution of intercepted photosynthetically active radiation (PAR). Shoots were collected from different heights in the crowns of trees representing four different size classes. A new method was developed to estimate seasonal light interceptance (SLI, intercepted PAR per unit needle area) of the shoots using a model for the directional distribution of above-canopy PAR, measurements of shoot silhouette area and canopy gap fraction in different directions. The ratio SLI/SLI(o), where the reference value SLI(o) represents the seasonal light interceptance of a spherical surface at the shoot location, was used to quantify the efficiency of light capture by a shoot. The ratio SLI/SLI(o) doubled from the top to the bottom of the canopy, mainly as a result of smaller internal shading in shade shoots than in sun shoots. Increased light-capturing efficiency of shade shoots implies that the difference in intercepted light by sun shoots versus shade shoots is much less than the decrease in available light from the upper to the lower canopy. For example, SLI of the five most sunlit shoots was only about 20 times greater than the SLI of the five most shaded shoots, whereas SLI(o) was 40 times greater for sun shoots than for shade shoots. Nitrogen content per unit needle area was about three times higher in sun needles than in shade needles. This variation, however, was not enough to produce proportionality between the amounts of nitrogen and intercepted PAR throughout the canopy.  相似文献   

3.
We assessed stem burial and adventitious root formation of two late-successional species, Abies mariesii and A. veitchii, in central Japan. In a plot (5 × 5 m), all seedlings between 8 and 24 cm tall were excavated: six A. mariesii seedlings in soil, and six and four A. veitchii seedlings in soil and on logs, respectively. For each sampled seedling, the number of terminal bud scars (TBS) was counted on the aboveground and belowground stems. Stem length was measured, and divided into aboveground and belowground stems. Among the three groups (A. mariesii seedlings in soil and A. veitchii seedlings in soil or on logs), there was no significant difference in height or total root weight (sum of adventitious roots and primary roots), but diameter at ground level and number of TBS were significantly different. Counting TBS on the aboveground stem of seedlings in soil underestimated seedling age, whereas the estimate was much closer to the true age for seedlings on logs. Seedlings in soil formed more adventitious roots than seedlings on logs. A large proportion of the stem was buried in humus for seedlings in soil, while most of the stem was not buried in humus for seedlings on logs. These results suggest that substrate affects adventitious root formation, the formation of which is important to shade tolerance. Thus, our preliminary results suggest that consideration of adventitious root formation is necessary to understand seedling bank dynamics and estimate seedling ages of these Abies species in spatially heterogeneous old-growth subalpine forests.  相似文献   

4.
Light-related plasticity in a variety of crown morphology and within-tree characteristics was examined in sun and shade saplings of Abies amabilis Dougl. ex J. Forbes growing in two late-successional forests with different snow regimes in the Cascade Mountains of Washington, USA. Compared with sun saplings, shade saplings typically had broad flat crowns as a result of acclimation at several scales (needle, shoot, branch, crown and whole sapling). Shoots of shade saplings had a smaller needle mass per unit of stem length than shoots of sun saplings, a feature that enhances light-interception efficiency by reducing among-needle shading. The low annual rate of needle production by shade saplings was associated with a longer needle lifespan and slower needle turnover. Reduced needle production within a shoot was reflected at the branch level, with lateral branches of shade saplings having a smaller needle mass than branches of the same length of sun saplings. Reduced allocation to needles permits greater investment in branches and stems, which is necessary to support the horizontally expanding branch system characteristic of shade saplings. Mean branch age of shade saplings was significantly higher than that of sun saplings. Shade saplings had lower needle mass per unit of trunk biomass or total biomass, reflecting greater investment in the trunk as a support organ. Increased investment in support organs in shade was more evident in the snowier habitat. The observed morphological acclimation makes A. amabilis highly shade and snow-tolerant and thus able to dominate in many late-successional forests in snowy coastal mountain regions.  相似文献   

5.
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   

6.
Structural characteristics of Abies mariesii M.T. Mast. saplings growing in sun and shade in a snowy subalpine parkland in central Japan were assessed to infer how saplings acclimate to suppression by larger individuals in a conifer clump and to extremely snowy conditions. Sun and shade saplings produced structurally different current-year shoots, and allocated biomass to needles and stem differently. Compared with sun saplings, shoots of shade saplings had lower needle mass per unit shoot size, which indicates less dense needle packing and more effective use of the limited available light by avoiding mutual shading among needles. Biomass allocation within lateral branches also differed between sun and shade saplings. Compared with sun saplings, needle mass was a smaller proportion of total branch mass in shade saplings although shade saplings retained needles for longer, thereby compensating, in part, for their lower annual production of needles. Thus shade saplings incur a high mechanical cost to support their low-light acclimated, conspicuously flattened crowns in this snowy habitat. Suppressed saplings are an important component of the persistent conifer clumps in snowy subalpine parklands. The observed structural characteristics of A. mariesii saplings, which ensure high shade- and snow-tolerance, contribute to the dominance of the species in snowy subalpine regions in Honshu, Japan.  相似文献   

7.
In 2010, the first, and so far only, infestation of the Hungarian spruce scale (Physokermes inopinatus) and accompanying sooty mould occurred in Scania, southernmost Sweden. About 1000?ha of Norway spruce (Picea abies) were affected, and the trees suffered from the sucking of the insects as well as from the dense sooty mould that covered the needles. Salvage cuttings were carried out in many of the massively attacked forest stands, both in response to the fear that the trees otherwise would die, e.g. from secondary bark beetle attacks and to prevent spreading of the infestation. The aim of this study was to provide basic, quantitative knowledge on the aftermath response of trees that were heavily infested, but not exposed to salvage cutting. Growth characteristics, in terms of needle weight, shoot length and tree-ring size were measured on infested and uninfested trees to compare and contrast the spruce growth before, during and after the scale outbreak. The infestation resulted in dwarf annual shoots, stunted needles and thin tree rings. The needle weight returned to normal the following year, whereas shoot length and tree rings required one growing season before full recovery.  相似文献   

8.
The ascomycete Nematostoma parasiticum (syn.: Herpotrichia parasitica) is commonly perceived as the causal agent of the so‐called Herpotrichia needle browning in silver fir (Abies alba). However, its fruitbodies are rarely present on symptomatic needles, which are also colonized by many presumably saprotrophic fungi. We compared the internal colonization of healthy and symptomatic needles on two sites in Poland. In addition, the endophytic mycobiota in needles of various age was recorded on two other sites without disease symptoms. Fungi were isolated from 95.6% of the dead needles and from 62.9% of the living needles on symptomatic trees, whereas on healthy trees, only 45.0% of the needles were colonized internally. Colonization frequency increased with needle age. From a total of 2017 isolates, 116 fungal taxa were identified. Frequency of many species was influenced by needle type. Anthostomella formosa, Gloeosporidiella sp., Hypoxylon fragiforme, Xylaria hypoxylon and X. polymorpha were the most common fungi isolated from living needles. In symptomless living needles, fungi occurred significantly more often in the basal than in the apical parts. In dead needles, the most common fungi were Alternaria alternata, Paraconiothyrium sporulosum, Fusarium sp., Mollisia cinerea, Rhizoctonia sp., Rhizosphaera oudemansii, Thysanophora penicillioides, Xylaria hypoxylon and X. polymorpha. Rhizoctonia sp. was the most frequently isolated fungus in dead needles (23.4%) but occurred rarely also in living needles (0.3–1.1%). The supposed pathogen N. parasiticum was detected only sporadically (at most in 0.6% of the needles). Our findings demonstrate the need for understanding the role of Rhizoctonia sp. in Herpotrichia needle browning disease aetiology.  相似文献   

9.
Total foliage dry mass and leaf area at the canopy hierarchical level of needle, shoot, branch and crown were measured in 48 trees harvested from a 14-year-old loblolly pine (Pinus taeda L.) plantation, six growing seasons after thinning and fertilization treatments.

In the unthinned treatment, upper crown needles were heavier and had more leaf area than lower crown needles. Branch- and crown-level leaf area of the thinned trees increased 91 and 109%, respectively, and whole-crown foliage biomass doubled. The increased crown leaf area was a result of more live branches and foliated shoots and larger branch sizes in the thinned treatment. Branch leaf area increased with increasing crown depth from the top to the mid-crown and decreased towards the base of the crown. Thinning stimulated foliage growth chiefly in the lower crown. At the same crown depth in the lower crown, branch leaf area was greater in the thinned treatment than in the unthinned treatment. Maximum leaf area per branch was located nearly 3–4 m below the top of the crown in the unthinned treatment and 4–5 m in the thinned treatment. Leaf area of the thinned-treatment trees increased 70% in the upper crown and 130% in the lower crown. Fertilization enhanced needle size and leaf area in the upper crown, but had no effect on leaf area and other variables at the shoot, branch and crown level. We conclude that the thinning-induced increase in light penetration within the canopy leads to increased branch size and crown leaf area. However, the branch and crown attributes have little response to fertilization and its interaction with thinning.  相似文献   


10.
The impacts of thinning, fertilization and crown position on seasonal growth of current-year shoots and foliage were studied in a 13-year-old loblolly pine (Pinus taeda L.) plantation in the sixth post-treatment year (1994). Length of new flushes, and their needle length, leaf area, and oven-dry weight were measured in the upper and lower crown from March through November. Total shoot length was the cumulative length of all flushes on a given shoot and total leaf area per shoot was the sum of leaf areas of the flushes.

By the end of June, first-flush foliage reached 70% of the November needle length (14.3 cm) and 65% of the final leaf area (15.0 cm2). Cumulative shoot length of first- and second-flush shoots achieved 95% of the annual length (30.3 cm), whereas total leaf area per shoot was 55% of the final value (75.3 dm2). Fertilization consistently stimulated fascicle needle length, dry weight, and leaf area in the upper crown. Mean leaf area of upper-crown shoots was increased by 64% six years after fertilization. A significant thinning effect was found to decrease mean leaf area per shoot in the crown. For most of the growing season, the thinned-fertilized trees produced substantially more leaf area per shoot throughout the crown than the thinned-nonfertilized trees. These thinned-fertilized trees also had greater needle length and dry weight, longer first flush shoots, and more leaf area per flush than trees in the thinned-nonfertilized plots. Needle length and leaf area of first flush shoots between April and July were linearly related to previous-month canopy air temperature (Ta). Total shoot length strongly depended on vertical light gradient (PPFD) within the canopy, whereas shoot leaf area was a function of both PPFD and Ta. Thus, trees produced larger and heavier fascicles, more and longer flush shoots, and more leaf area per shoot in the upper crown than the lower crown. We conclude that thinning, fertilization, and crown position regulate annual leaf area production of current-year shoots largely by affecting the expansion of first flush shoots and their foliage during the first half of the growing season.  相似文献   


11.
The effects of decapitation at various levels, combined with heavy pruning of remaining branches, were examined for Caribbean pine. This treatment stimulated the development of shoots with long primary needles at all levels in the crown of trees aged one, four and eight years. Although a significant positive relationship between primary needle length and rooting was demonstrated, the very juvenile appearance of these shoots was not a reliable guide to rooting. There was an over-riding effect on rooting of the ortet age × level in crown × decapitation height treatment. Decapitated trees provided shoots of higher rooting capacity than intact trees for the one and four, but not the eight year old trees. Shoots developing in the lower crown in response to decapitation and heavy pruning of four year old trees displayed a high level of rooting-equivalent to that of shoots from the decapitated and pruned one year old trees.  相似文献   

12.
We compared the range and variation in shoot silhouette area to projected leaf area ratio (SPAR) in fertilized and unfertilized (control) Norway spruce (Picea abies (L.) Karst.) trees. We measured SPAR for several view directions of 169 shoots at different depths in the crown of fertilized and control trees. There was an increase in SPAR with depth in the crown in both control and fertilized trees. In the fertilized trees, however, mean SPAR was larger overall, the increase with depth in the crown was steeper, and there was a larger variation in SPAR with inclination and rotation angle of the shoot (relative to the view direction). In particular, shoots in the lower crown of fertilized trees were rotationally asymmetrical ("flat") and had high values of the maximum ratio of shoot silhouette area to projected leaf area (SPAR(max)). Differences in SPAR between fertilized and control trees were explained by changes in shoot structure in response to fertilization and shading. Shoots of fertilized trees were larger and had more needle area than shoots of control trees. However, the ratio of needle area to shoot size was smaller in fertilized trees than in control trees, implying less within-shoot shading and, consequently, a larger SPAR. Also, the increase in SPAR with increased shading (depth in the crown) could be explained by a decrease in the ratio of needle area to shoot size. In addition, because fertilized trees had more needle area than control trees, the effect of shading at a given depth in the crown was more pronounced in fertilized trees than in control trees.  相似文献   

13.
The effect of two training systems (Central Leader with branch pruning versus Centrifugal Training with minimal pruning, i.e., removal of fruiting laterals only) on canopy structure and light interception was analyzed in three architecturally contrasting apple (Malus domestica Borkh.) cultivars: 'Scarletspur Delicious' (Type II); 'Golden Delicious' (Type III); and 'Granny Smith' (Type IV). Trees were 3D-digitized at the shoot scale at the 2004 and 2005 harvests. Shoots were separated according to length (short versus long) and type (fruiting versus vegetative). Leaf area density (LAD) and its relative variance (xi), total leaf area (TLA) and crown volume (V) varied consistently with cultivar. 'Scarletspur Delicious' had higher LAD and xi and lower TLA and V compared with the other cultivars with more open canopies. At the whole-tree scale, training had no effect on structure and light interception parameters (silhouette to total area ratio, STAR; projected leaf area, PLA). At the shoot scale, Centrifugal Training increased STAR values compared with Central Leader. In both training systems, vegetative shoots had higher STAR values than fruiting shoots. However, vegetative and fruiting shoots had similar TLA and PLA in Centrifugal Trained trees, whereas vegetative shoots had higher TLA and PLA than fruiting shoots in Central Leader trees. This unbalanced distribution of leaf area and light interception between shoot types in Central Leader trees partly resulted from the high proportion of long vegetative shoots that developed from latent buds. These shoots developed in the interior shaded zone of the canopy and therefore had low STAR and PLA. In conclusion, training may greatly affect the development and spatial positioning of shoots, which in turn significantly affects light interception by fruiting shoots.  相似文献   

14.
We tested the effects of thinning on allometry and needle-age distribution in natural stands of Abies sachalinensis Masters by comparing a thinned stand to an unthinned, control stand. Specifically, we attempted to clarify how allometry was altered after a thinning. We assumed that the needle-age distribution of trees in the thinned stand would show a younger composition than in the control stand, given the effect of improved light conditions on needle dynamics following a thinning. These investigations were conducted in dense Abies stands located in central Hokkaido, northern Japan, 19 years post-thinning. In the thinned stand, the ratio of individual needle mass to stem mass increased significantly, as compared to the control. A difference in the H–DBH relationship between the stands was probably related to this tendency. Mean needle age of trees differed significantly between the two stands, and the thinned stand showed a younger needle age than the control. Within each stand, dominant trees showed older mean needle age than codominant or suppressed trees. These tendencies may have been caused by differential needle dynamics affected by light conditions in the stands, and by different crown positions among the trees within a stand. In summary, trees in the thinned stand showed increased growth rates after thinning, which were caused by increased needle mass, younger composition of needles, and improved light conditions.  相似文献   

15.
J. Fuhrer 《Forest Pathology》1985,15(4):227-236
Measurements of ethylene production, free and conjugated ACC concentrations, chlorophyll content, and length of needles from fir trees (Abies alba Mill.) growing in stands affected by “Waldsterben” showed that during the development of the disease two consecutive stages can be distinguished: a. long-term, low ethylene production and accumulation of conjugated ACC (MACC) in response to chronic stress, associated with reduced needle growth, and b. high ethylene production associated with chlorophyll breakdown, followed by premature abscission of the needles.  相似文献   

16.
We investigated effects of nutrient availability on shoot structure and light-interception efficiency based on data from control (C) and irrigated + fertilized (IL) trees of Norway spruce (Picea abies (L.) Karst.). The sampling of 1-year-old shoots was designed to cover the variation in canopy exposure within the live crown zone, where current-year shoots were still found. Canopy openness was used as a measure of light availability at the shoot's position. Openness values for the sample shoots ranged from 0.02 to 0.77 on the IL plot, and from 0.10 to 0.96 on the C plot. Among needle dimensions, needle width increased most with canopy openness. At fixed canopy openness, needle width was larger, and the ratio of needle thickness to width was smaller in IL trees than in C trees. Specific needle area (SNA) and the ratio of shoot silhouette area to total needle area (STAR) decreased with canopy openness, so that the combined effect was a threefold decrease in the ratio of shoot silhouette area to unit dry mass (SMR = STAR x SNA) along the studied range of openness values. This means that the light-interception efficiency of shoots per unit needle dry mass was three times higher for the most shaded shoots than for sun shoots. A test of the effect of fertilization on the relationships of SNA, STAR and SMR indicated statistically significant differences in both slope and intercept for SNA and STAR, and in the intercept for SMR. However, the differences partly cancelled each other so that, at medium values of canopy openness, differences between treatments in predicted SNA, STAR and SMR were small. At 0.5 canopy openness, predicted STAR of IL shoots was 6.1% larger than STAR of C shoots, but SMR of IL shoots was 10% smaller than that of C shoots. The results suggest that light-interception efficiency per unit needle area or mass of the shoots is not greatly affected by fertilization.  相似文献   

17.
Pensa M  Jalkanen R 《Tree physiology》2005,25(10):1265-1271
Latitudinal variation in needle longevity of conifers in response to climatic variability is a well-known phenomenon, but its significance has only rarely been studied. Scots pine (Pinus sylvestris L.) stands were investigated in four locations in Lapland (northern Finland and northern Sweden, 67-68 degrees N) and in four locations in Estonia (59 degrees N) to test the hypothesis that plasticity in needle longevity results in similar needle biomass per foliated shoot length across latitudes. The needle-trace method (NTM) revealed the dynamics of needle fascicles attached to stem shoots. Long-term mean needle age was 2 years greater in Lapland than in Estonia. In both regions, mean needle age was strongly and negatively related to yearly needle-fascicle production rate relative to the number of needles on the same shoot axis. Although significantly fewer needle-fascicles were produced annually in trees in Lapland than in Estonia, the overall number of fascicles attached to the stem shoots, needle-fascicle density and individual needle dry mass did not differ between regions. Consequently, needle biomass per foliated shoot length was similar in trees in both regions. Thus, our results support the theory that plasticity in needle longevity helps Scots pine to compensate for reduced needle production so that the same foliar biomass per shoot is retained under a wide range of growth conditions.  相似文献   

18.
The patterns of current‐year shoot, needle and terminal bud elongation in seedlings of three Scots pine (Pinus sylvestris L.) and three lodgepole pine (Pinus contorta Dougl. var. latifolia Engelm.) provenances were compared during the third and fourth growing seasons after planting. Lodgepole pine produced longer shoots and buds than did Scots pine, mainly because lodgepole pine formed more stem units and elongated at a faster rate. Stem unit length and the duration of shoot and bud elongation differed relatively little between species and provenances. Lammas or polycyclic growth occurred in some lodgepole pine provenances, but not in any Scots pine provenance, and was associated with enhanced shoot elongation. Needle elongation commenced earlier, proceeded at a faster rate, and was greater in lodgepole pine than in Scots pine, but ceased about the same time in all species and provenances. The heat sum required to attain 50% of final length was lower for shoots and needles in lodgepole pine than in Scots pine, and for shoots in northern provenances than in southern ones. Mitotic activity in the apical meristem of the terminal bud, which occurred less than one week after the seedlings were free from snow, started and ceased about the same time in each species, but was higher in lodgepole pine than in Scots pine early in the shoot elongation period.  相似文献   

19.
The number of short shoots per shoot length, or needle density, is species typical, and it shows year‐to‐year variation within species. By modification of the needle trace method, long‐term needle density chronology was produced in a Scots pine (Pinus sylvestris L.) stand, located at the northern timberline in Finland. Treewise, needle density varied between 9 and 14 short shoots per long shoot (stem internode) centimetre, the annual minimum and maximum values being 5 and 37 short shoots cm?1. The stand‐specific long‐term average was 10.5 short shoots cm?1, and the mean annual value varied between 17 and 8 short shoots cm?1 in 1951 and 1984, respectively. The long‐term pattern in needle density was one of decline with time between 1950 and the mid‐1970s, then to slightly increase on entering the 1990s. The years when the density was relatively high were 1957, 1968 and 1981, indicative of some climatic extremes.  相似文献   

20.
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   

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