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1.

Context

The knowledge on cambial activity in water-limited environments, such as the Mediterranean, is still fragmentary. Dendrochronological studies have determined that spring precipitation plays an important part in determining tree-ring width and the properties of tracheids. However, the complex relation between cambial phenology and climate is still far from understood.

Aims

We studied the influence of climate, especially water stress, on maritime pine wood formation with the aim of determining the influence of drought on cambial activity.

Methods

A plantation of maritime pine (Pinus pinaster) was selected in the west coast of Portugal, to monitor cambial activity and wood formation using anatomical observations and band dendrometers. The trees were monitored weekly over 2 years (2010 and 2011).

Results

Xylem differentiation started earlier when warmer late winter temperatures were observed. Water stress triggered an earlier stop of wood formation and also the formation of tracheids with smaller lumen area. In both years a bimodal pattern of stem radial increment was registered by band dendrometers with two periods of increment: one in spring and another in autumn. The xylem anatomy study suggests that the autumnal increment period corresponded mostly to stem rehydration, since the differentiation of new xylem cells by the cambium was not observed.

Conclusion

Maritime pine cambial activity appears to be under a double climatic control: temperature influences cambial onset and water availability growth cessation.  相似文献   

2.
Examining ecological limits to shade acclimation at whole-plant level is determinant for evaluating the success of sapling establishment in low-light environments. We studied nutritional effects on whole-plant development in response to shade in two Mediterranean forest tree species with different successional status: the early-successional Pinus halepensis Mill. and the late-successional Quercus pubescens Wild. Through a nursery-based factorial experimental design approach, we measured height increment along 2 years and final leaf, stem and root biomass in both species saplings subjected to two lights and two soil nutrient availability treatments. The shade avoidance response was exclusive to P. halepensis, appeared as timely dependent, and persisted longer in saplings exposed to higher nutrient availability. Q. pubescens benefited from the higher nutrient availability by lowering the light-driven plastic response in aerial support investment and belowground carbon allocation, whereas P. halepensis heightened its light-driven plastic response. These contrasted responses are thus clearly related to the shade acclimation strategy of each species: the shade-intolerant P. halepensis enhances shade avoidance when non-nutrient-limited, whereas the shade-tolerant Q. pubescens assumes a conservative strategy by limiting phenotypic plasticity-induced costs. Maintaining greater shade avoidance in non-nutrient-limited soil conditions might be an adaptive advantage for P. halepensis seedlings growing in the understory, in response to gap formation in the overstory. In contrast, the more conservative and less costly shade responsiveness of Q. pubescens may confer it a better adaptive advantage in long-term light-limited environments.  相似文献   

3.
The Mediterranean evergreen woody plants Quercus coccifera and Pinus halepensis grow in a range of environments where selection by drought, heat and high irradiance can drive genetic and phenotypic differentiation of populations. However, the role of these stresses in filtering out maladaptive genotypes remains unknown. We hypothesize that this filtering is an important process for woody Mediterranean species due to their low phenotypic plasticity reported in previous studies. We have studied the response of saplings of Q. coccifera and P. halepensis, originating from two contrasting populations (a rock outcrop and a garrigue formation), to water stress. Isozyme characterization of genetic diversity was done to determine whether populations were genetically distinct. Water response analysis was based on water relations, gas exchange, chlorophyll a fluorescence, pigment content, antioxidant status and morphological and structural parameters. Ecotypic differentiation was found for both Q. coccifera and P. halepensis populations, with a higher population isozyme similarity and a higher frequency of dominance of a few genotypes at the rock outcrop in both the species. P. halepensis exhibited small but significant differences between populations for plastic responses to water, with lower phenotypic plasticity in saplings from the rock outcrop. Although it was not found in Q. coccifera, this pattern suggests that ecotypic differentiation rendering stress-tolerant ecotypes involves a decreased plasticity. Phenotypic plasticity was not high but it explained over 75% of the total variability among individual plants. Thus, and although evidence for ecotypic divergence was found in both the species, saplings were plastic enough to blur ecotypic differentiation. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.  相似文献   

4.
Purportedly, large Douglas-fir trees in the American Pacific Northwest use water stored in bole tissues to ameliorate the effects of seasonal summer drought, the water content of bole tissues being drawn down over the summer months and replenished during the winter. Continuous monitoring of bole relative water content (RWC) in two 110-120-year-old Douglas-fir trees with ThetaProbe impedance devices provided an integrated measure of phloem-sapwood water content over 4 years. Seasonal changes in RWC closely tracked cambial activity and wood formation, but lagged changes in soil water content by 2-3 months. The RWC in the combined phloem and sapwood markedly increased during earlywood production in the late spring and early summer to maximum values of 64-77% as plant available soil water (ASW) decreased by approximately 60%. With transition and latewood formation, RWC decreased to minimum values of 59-72%, even as ASW increased in the fall. The difference between minimum RWC in the fall and maximum RWC in midsummer was only approximately 5%. Seasonal changes in bole RWC corresponded to cambial phenology, although decreasing AWS appeared to trigger the shift from earlywood to latewood formation.  相似文献   

5.
Xylogenesis was monitored during 2003 and 2004 in a timberline environment in southern Italy to assess links between temperature, cambial phenology and wood formation on a short-time scale. Wood microcores were collected weekly from May to October from 10 trees of Pinus leucodermis Ant., histological sections were cut with a rotary microtome and anatomical features of the developing and mature tracheids were observed and measured along the growing tree ring. Spring 2003 was hotter than spring 2004, with temperatures up to 2.6 degrees C above historical means. The hotter conditions resulted in an earlier onset of cambial activity and all differentiation phases of about 20 days, resulting in an increased duration of xylogenesis of about 23 days. Air and stem temperatures at which xylogenesis had a 0.5 probability of being active were calculated with logistic regressions fitted on binary responses. In both years, similar thresholds were estimated with daily mean values of 8.2 and 9.5 degrees C for air and stem temperatures, respectively. The observed convergent responses of cambium phenology to temperature during the two contrasting springs confirm the key role of this environmental factor in determining the onset and duration of wood formation in timberline areas. The intra-annual dynamics of ring-width increase differed between years, with significantly narrower rings formed in 2004 than in 2003. These differences were mainly related to cell size because larger earlywood tracheids were produced in 2003. This study demonstrates the plasticity of tree-ring formation in response to high temperatures as a result of modifications in the onset and duration of differentiation.  相似文献   

6.
Pinus halepensis has been described as a drought-tolerant species with high plasticity to growth in different environments. Its eco-physiological characteristics could facilitate the use of this species in large afforestations in the future scenery of climate change. Somatic embryogenesis is a biotechnological tool with potential for large-scale clonal propagation. In order to establish an improved regeneration protocol for Pinus halepensis, the effects of different temperatures (18, 23, and 28 °C) and water availability conditions (2, 3, and 4 g L?1 Gelrite®), during initiation of embryonal masses on the rate of initiation, proliferation, maturation, and the number of embryos developed, were evaluated. It was found that environmental conditions during the initiation stage of Pinus halepensis somatic embryogenesis influence the success of initiation and proliferation. In contrast, there was no effect of these conditions on the maturation rates and the number of somatic embryos. Somatic embryos were obtained in all treatments tested, indicating that plants can be produced from extreme conditions of induction, such as high temperatures (28 °C) and low water availability conditions (4 g L?1).  相似文献   

7.
The purpose of the present study is to model the regeneration success using quantifiable growth factors. The hypothesis is that the regeneration success can be predicted from a model that incorporates canopy and ground flora leaf area indexes (LAI), soil water content and soil carbon content. In April 1992, 10 plots (1 m2) were established in each of 22 Danish beech stands with natural regeneration originating from the 1989 seed fall. The regeneration success was investigated until autumn 1994. In each of the 220 plots the light conditions, soil water supply and soil carbon content were quantified. The survival was high during the study period. From spring 1992 to autumn 1993 the mean number of saplings decreased from 58 m−2 to 54 m−2. The sapling number was reduced by increased canopy density but it was neither influenced by water supply nor by soil carbon content. The height growth variation was evident. After the fifth growing season (autumn 1994) the height of the tallest sam3pling per plot ranged from 7 to 120 cm. Generally, an increased canopy opening increased the potential for and variance of height growth. Increased soil water content caused a significant increase in the growth of the saplings; whereas, a significant reduction in growth occurred with increased soil carbon content. The shading from the competing ground flora had no significant effect. However, the contribution to the total sum of squares from the quantitative variables of the relevant statistical models was only 3%–13%. Therefore, the models were unable to predict regeneration success satisfactorily and it was not possible to establish the causal effects underlying silvicultural treatments and site. It is concluded that the beech saplings probably were so well established at the beginning of the study that they had become tolerant to poor growing conditions, such as the drought in 1992. This indicates that the growing conditions during the early phases of seedling establishment may be crucial for the regeneration success. For silviculturists, these conclusions stress the importance of creating favourable growth conditions prior to the first growing season. Future research should examine the early phases of seedling establishment by quantifying growth factors in more detail than was done in the present study.  相似文献   

8.
To assess and improve methods for the development of coastal forests in Hokkaido, northern Japan, this study examined the factors affecting mortality of planted Pinus thunbergii Parlat. saplings in a region with seasonal soil frost. The needles of pine saplings at the study site turn red in late spring, suggesting xylem embolism, which can lead to death. Sapling mortality was strongly correlated with the degree of cold just after planting and with the occurrence of repeated severe winters. Volcanic ash soil had been supplied to improve the soil nutrients, and the volcanic ash soil always thawed later than the original coastal sand. Saplings in volcanic ash were significantly more damaged than those in original coastal sand. This suggests that hydraulic stress caused by cuticular transpiration under frozen soil conditions during spring may also accelerate damage due to embolisms. Hence, volcanic ash soil should not be applied to soils in regions with seasonal soil frost. A covering of woodchips can help prevent frost from reaching greater soil depths, and, indeed, fewer saplings at the study site died in areas with such coverage. Planting location can also affect pine sapling mortality. For example, shading caused by wood fencing may affect the soil freezing profile during winter; therefore, planting close to a fence should also be avoided in regions with seasonal soil frost. These findings should be used to improve the development of forests in regions with seasonal soil frost.  相似文献   

9.
油松管胞形态特征的变异   总被引:13,自引:0,他引:13  
徐有明 《林业科学》1990,26(4):337-343
本文研究分析了山西中条山产地油松管胞形态特征的变异。管胞长度自髓心向外,首先迅速增加,13年后管胞长度增加缓慢,20年后保持相对稳定。管胞长度沿树干主轴自基部向上逐渐增加,5.3m高处最长,然后向上变短,树冠区域管胞长度最短。形成层原始细胞长度随着原始细胞年增大,开始递增,达到最大值后又递减。管胞直径、胞壁厚度自髓心向外增加。管胞直径轴向变化由树干基部开始向上增大,然后又减小。管胞长宽比、壁腔比的径向变异与管胞长度的径向变异模式相似。生长轮内管胞长度从早材到晚材,开始减小,然后增加,最小值位于早晚材过渡处。  相似文献   

10.
To determine the effect of burnt tree removal on post-fire natural regeneration of Pinus halepensis, two 2 500 m2 areas were selected six months after the fire in a totally destroyed mature (>70 years) pine forest. In one area, all the trees were cut down and removed 10 months after the fire and, in the other, all the trees were left standing (control). In each area, 20 permanent plots of 20 m2 each were randomly placed, and all seedlings emerging within them labeled by individual numbered plastic tags. Emergence, mortality, density and growth (height) of 6649 P. halepensis seedlings were monitored during the first four post-fire years. Seedling emergence was concentrated in the first post-fire autumn–winter period. No positive effect on seedling emergence was detected as a consequence of burnt trunk dragging and subsequent turning over of soil. Wood removal produced an immediate average seedling mortality of 33%, and notably increased seedling mortality during the subsequent summer, probably due to increased exposure of seedlings to sunlight and the possible debilitation of many individuals by mechanical contact during burnt wood removal. A negative correlation of pine seedling mortality with height was detected, which increased significantly on wood removal in the third post-fire year. That is, short seedlings (<10 cm) in treated plots were the most likely to die during this period. In spite of the detrimental effect of wood removal on sapling survival, seedling density four years after fire in the cleared area was large (3.3 seedlings/m2). Wood removal treatment reduced seedling growth: seedling height was significantly higher in the control stand, and differences in seedling height growth rate became particularly noticeable in the fourth post-fire year. The results denote that traditional wood removal practices do not threaten natural post-fire P. halepensis re-establishment if initial seedling density is large enough. However, further studies focused on wood removal effects on the final tree development level and other ecological aspects are necessary to choose adequate post-fire forest management.  相似文献   

11.
12.
The effects of removing aboveground vegetation on survival and growth of Hondo spruce (Picea jezoensis var. hondoensis) saplings were examined in a dense dwarf bamboo (Sasa nipponica) field inside a deer-proof fence built in a degraded sub-alpine forest on Mt Ohdaigahara, central Japan. All bamboo culms were removed within a radius of 1 m from each sapling in a plot of 0.6 ha (removal plot), whereas no culms were removed in two control plots of 0.1 ha. We measured the height, crown depth, and main shoot elongation over 1 year for all spruce saplings, culm height and cover ratios of dwarf bamboo growing around each spruce sapling, and the light levels above each sapling. Removal of dwarf bamboo regulated the culm height and bamboo coverage until the following summer such that light conditions improved. The mortality of saplings smaller than the average culm height was lower in the removal plot (0.89%) than in a control plot (5.9%). After bamboo removal, sapling height growth was reduced with increasing bamboo cover in the previous year. Complete removal of the aboveground portions of covering bamboo improved sapling survival while regulating elongation growth, possibly because of the sudden increase in light intensity. Thus, controlling the extent of dwarf bamboo removal might be necessary to facilitate the growth of spruce saplings to restore the forest.  相似文献   

13.
Three-year-old Alnus glutinosa (L.) Gaertn. (alder) saplings were single or double inoculated at the stem base with Phytophthora alni subsp. alni Brasier & S.A. Kirk under natural climatic conditions. Lesion formation on the bark showed a biphasic pattern of development, with extension occurring at a moderate rate in spring, and more rapidly during late summer. However, large variability was encountered in pathogen development within the population of infected saplings, ranging from high susceptibility to almost complete resistance. Infection resulted in severe growth retardation, and death within two years of inoculation in 75% of the saplings. During disease development, rates of transpiration and CO(2) uptake were significantly reduced. Consequently, minimum leaf water potentials were less negative in infected saplings than in control saplings. Surviving saplings matched control trees in photosynthetic capacity, transpiration rate and water potential during the second year of infection. Leaf starch concentration of infected saplings was significantly higher than in control saplings, possibly indicating that the destruction of bark tissue by the pathogen impaired phloem transport from leaves to roots.  相似文献   

14.
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   

15.
A better understanding of the ecophysiological basis of wood formation by monitoring radial growth over the whole vegetation period may help to explain possible discrepancies between long-term average climate–growth relationships and short-term climatic impacts on tree growth. To understand how growth–climate associations of Oriental beech vary throughout a vegetation period, we studied seasonal growth patterns of high-elevation beech trees growing in the north of Iran by collecting wood anatomical micro-cores in 10 to bi-weekly intervals and measuring stem increment with high-resolution electronic dendrometers. Wood formation was for two consecutive years with contrasting inter-annual climate conditions (2011 and 2012). We divided the growing period into three equal time phases and related daily climate variability to the cambial growth in each phase. The pattern of climate–growth relationships varies over a complete growing season and between years: in both study years, trees responded homogenously at the beginning and at the end of the growing season, but showed opposing influence of relative humidity during spring and early summer (June and July). Temperature as the main driver of xylogenesis had a stimulating effect on growth at the beginning of the growing season, but had negative effects on radial increment during late June and July mainly due to the excess of ambient temperatures over an optimum threshold. Higher temperature in late June and July 2011 compared with those of 2012 led to a significantly narrower tree ring in 2011, despite the similar sums/means of annual climate parameters in these two years. Since regional climate change scenarios expect higher temperature and reduced moisture conditions in future, radial growth of beech in the north of Iran may be adversely affected.  相似文献   

16.
Marsal J  Girona J 《Tree physiology》1997,17(5):327-333
Effects of water deficits on leaf turgor maintenance processes were analyzed for pear trees (Pyrus communis L. cv. "Barlett") grown in 120-liter containers. Four irrigation treatments were applied: a well-watered control treatment, a spring water stress cycle (Sp), a summer water stress cycle (Su), and a spring plus summer water stress cycle (Sp + Su). For the Sp treatment, water application was progressively reduced from 100 to 20% of the control dose over a period of 27 days in spring. For the Su treatment, water application was progressively reduced over 23 days in summer, from 100 to 20% of the control dose. The Sp + Su treatment comprised both the spring and summer drought stress cycles. Pressure-volume (P-V) curves were constructed and stomatal conductances were determined for pear leaves from each treatment during the spring and summer stress cycles. Leaf water potential (Psi(pi) (0)) and relative water content (R(0)) at the turgor loss point of control leaves tended to decrease from spring to summer. Changes in leaf osmotic water potential at full turgor (Psi(pi) (100)) and in symplast water fraction (R(s)) did not explain the seasonal decrease in Psi(pi) (0). The water stress treatments had no effect on Psi(pi) (100), but R(s) was reduced by the water stress treatments, particularly during the summer stress cycle of the Su and Sp + Su treatments. The decrease in R(s) was correlated with an increase in the slope of the linear region of the P-V curve. Such a coupled adjustment would lead to increased water uptake capacity of water-stressed trees only under non-turgor conditions. Furthermore, pear leaves did not actively accumulate solutes. We conclude, therefore, that changes in leaf tissue water relations as a result of leaf acclimation to water stress are unlikely to facilitate maintenance of fruit productivity under drought.  相似文献   

17.
Neighboring trees growing under identical environmental conditions can exhibit different dynamics and periods of growth. Despite the recent advances in cambial biology, the exogenous and endogenous factors generating asynchronous xylem growths still remain undetermined. This study investigated timings and duration of xylem formation in maritime pine (Pinus pinaster Ait.) from an even-aged plantation in Portugal growing under Mediterranean climate. Cambial phenology and stem diameter were monitored weekly, from March to December 2010, on two classes of trees divided according to the tree ring widths of the last 15 years, but similar age and size: fast- and slow-growing trees. We tested the hypothesis that differences in tree ring widths result from cell production which in turn affects timings of xylogenesis and that the bimodal growth pattern, typical of the Mediterranean, originates from a double reactivation of the cambium: in spring and autumn. Cambial activity started earlier and ended later in fast-growing trees, confirming that cell production is a key factor determining the duration of xylogenesis. Intra-annual variations in stem diameter recorded by band dendrometers revealed two peaks of increment occurring in spring and late summer. However, the number of cambial cells did not increase in late summer, which suggested that the second peak of increment was caused by stem rehydration, rather than by a reactivation of cell division. These results demonstrated that the variability in the timings of xylem phenology observed among trees of the same age and size and growing under similar environmental conditions was closely related to cell production and not to age or size per se.  相似文献   

18.
Effects of the timing of soil thawing in the spring on Scots pine (Pinus sylvestris L.) were studied under controlled laboratory conditions. Sixteen 6-year-old saplings were lifted from the field, replanted in containers and placed in four treatments in controlled environment (CE) chambers with four replicate saplings per chamber. The saplings were held in the CE chambers during one simulated winter and one simulated growing season. The soil was frozen to -2 degrees C during a second simulated winter in the CE chambers, and the soil thawing treatments began at the end of the second simulated winter. Soil thawing began at various times before (no delay in thawing) and after (delay in thawing) chamber air conditions were changed from simulated winter to simulated summer. Delayed soil thawing subjected saplings to stress, with the severity of stress depending on the length of the delay in thawing. If there was no delay or only a short delay in soil thawing, stress was minor and reversible. A 2-week delay in soil thawing led to death of the saplings. Stress was apparent as decreases in the variable to maximal chlorophyll fluorescence ratio (Fv/Fm), chlorophyll a/b ratio and needle water potential. In needles of stressed saplings, apoplastic electrical resistance first decreased and then increased and there were anomalies in the electrical impedance spectra of the stems. Stress from the soil thawing treatments affected both root and shoot growth.  相似文献   

19.
We monitored leaf phenology and xylogenesis of 12 Pedunculate oaks in northern Poland in 2014. We hypothesized that the individual trees, which differed in size, age and habitat (tree stand or gap), also diverged in terms of the seasonal patterns of leaf phenology and xylogenesis. The samples used for wood formation observations were collected most frequently during the early leaf phenophases (from March to end of June). The transverse sections of the cambial region were cut with a sledge microtome. We counted the number of cambial cells, measured the width of xylem increment and assessed the timing of xylogenesis and earlywood-vessel formation. We found significant differences in leaf phenology and timing of xylogenesis among individual trees. The smallest differences in wood formation among the trees were observed at the beginning of the vegetation season when the first earlywood vessels were detected (9 days). The dates of completion of the first tangential row of earlywood vessels varied by up to 30 days, while for the completion of the entire earlywood dates varied by up to 32 days. The highest productivity of cambial cells (13 cell layers) was observed around the time of bud swelling at mid-April. In the last days of April, the number of cambial cell layers decreased and subsequently increased again when the leaves were nearly fully expanded at the end of May. To summarize, we observed a high seasonal variability in the number of cambial cell layers. Differences in the time of cessation of cambial activity and xylogenesis amounted to 1 month. We conclude that: (1) oak tree-ring widths and earlywood-vessel sizes and numbers may not be sensitive indicators for early spring temperature and spring defoliation; (2) the missing association between leaf phenophases and xylogenesis as well as the phenological variability may be the reasons for the lack of a clear climatic effect on the above-mentioned parameters.  相似文献   

20.
应用定量解剖学和显微图像分析技术 ,得到了毛白杨枝条形成层细胞与其衍生木质部细胞解剖特征在形成层活动期内的变化情况 ,并探讨了两者之间的关系。采用X射线微密度测定技术 ,了解了同一生长轮内木材密度沿径向方向上的变异 ,讨论了其与木质部细胞解剖特征之间的关系。结果表明 ,形成层纺锤形细胞的径向和弦向宽度在活动期不同阶段差异不显著 ,而纤维长度、纤维宽度、壁腔比、微纤丝角、导管分布频率和胞壁率差异极显著 ;导管组织比量差异显著 ;导管分子长度、纤维比量和射线比量差异不显著。同一生长轮内木材密度的径向变异与活动期不同阶段形成的木质部细胞数量及其解剖特征直接相关。形成层带细胞层数与纤维长度、纤维长宽比和壁腔比呈极显著负相关 ,与纤维宽度呈显著正相关 ;形成层纺锤形细胞的径向宽度与纤维长度、纤维长宽比和射线比量呈显著负相关 ;形成层纺锤形细胞的弦向宽度与纤维宽度呈显著正相关。总体看来 ,在整个形成层活动期 ,形成层纺锤形细胞与木纤维的解剖特征之间随时间变化的相关性较好 ,而与导管分子的相关关系则不明显  相似文献   

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