Seasonal dynamics of wood formation: a comparison between pinning,microcoring and dendrometer measurements

Three different methods were evaluated for analysing wood formation of Norway spruce [Picea abies (L.) Karst.] and Scots pine (Pinus sylvestris L.) in Finland. During two growing seasons, wood formation dynamics were determined both by wounding the cambium with a needle followed by localisation of the wound-associated tissue modification after the growing season (pinning), and by extracting small increment cores during the growing season (microcoring). Stem radius was additionally monitored with band dendrometers. For Norway spruce, pinning and microcoring yielded similar dates for the onset of wood formation. The timing of wood production during the growing season was also similar for pinning and microcoring. For Scots pine, the onset of wood formation was recorded from microcores almost 2 weeks later than from pinning samples. In Scots pine, microcore measurements also produced somewhat later cessation dates for tracheid formation than the pinning samples. For both tree species, the total number of tracheids formed during the growing season was, however, about the same for pinning and microcoring. Dendrometer results clearly differed from those of pinning and microcoring. In particular, the dendrometers showed an increase of stem radius considerably earlier in spring, when the other methods did not detect wood formation. Thus, pinning and microcoring currently represent the most reliable techniques for detailed monitoring of wood formation.     

Effects of climate variables on intra-annual stem radial increment in Pinus cembra (L.) along the alpine treeline ecotone

? Within the alpine treeline ecotone tree growth is increasingly restricted by extreme climate conditions. Although intra-annual stem growth recorded by dendrometers can be linked to climate, stem diameter increments in slow-growing subalpine trees are masked by changes in tree water status.

? We tested the hypothesis that intra-annual radial stem growth in Pinus cembra is influenced by different climate variables along the treeline ecotone in the Austrian Alps. Dendrometer traces were compared with dynamics of xylem cell development to date onset of cambial activity and radial stem growth in spring.

? Daily fluctuations in stem radius reflected changes in tree water status throughout the treeline ecotone. Extracted daily radial increments were significantly correlated with air temperature at the timberline and treeline only, where budburst, cambial activity and enlargement of first tracheids also occurred quite similarly. A close relationship was detected between radial increment and number of enlarging tracheids throughout the treeline ecotone.

? We conclude that (i) the relationship between climate and radial stem growth within the treeline ecotone is dependent on a close coupling to atmospheric climate conditions and (ii) initiation of cambial activity and radial growth in spring can be distinguished from stem re-hydration by histological analysis.

     

Xylogenesis of Pinus pinaster under a Mediterranean climate

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.     

Daily patterns of stem size variation in irrigated and unirrigated Eucalyptus globulus

High resolution measurements of stem diameter variation provide a means to study short-term dynamics of tree growth and water status. In this 14-month study, daily changes in stem radius of Eucalyptus globulus Labill. seedlings were measured with electronic point dendrometers in a plantation in southern Tasmania, Australia. The daily patterns of stem expansion and shrinkage were classified into three phases: shrinkage; recovery; and increase in diameter from one maximum to the next, or increment. This study showed that rapid onset of even mild drought in irrigated trees caused distinct changes in daily patterns of stem diameter variation, particularly the duration of daily stem increment. The duration of the daily increment phase was directly related to increment magnitude. The dynamics of daily increment were significantly affected by mean minimum temperature, indicating a temperature limitation on metabolic processes underlying diameter growth in these trees. Most likely due to differences in conductance, the duration but not rate of the incremental daily expansion was greater in fast- than in slow-growing trees.     

Effect of flooding of soil on growth,stem anatomy,and ethylene production of cryptomeria japonica seedlings

Flooding of soil for 55 days altered the rate of growth and stem anatomy of 9‐month‐old Cryptomeria japonica seedlings. Although flooding did not affect height growth it reduced the rate of dry weight increment of seedlings while increasing stem diameter. The reduction in dry weight increment of seedlings resulted largely from decay of roots and, to a lesser extent, from inhibition of growth of roots and needles. The increased diameter growth of flooded seedlings resulted largely from an increase in bark thickness associated with increased phloem production and greater amount of intercellular space. Flooding reduced xylem increment in submerged stems but increased it above the water level because of larger tracheids rather than more tracheids per radial file. Flooding also increased lumen diameters of tracheids, decreased tracheid wall thickness (as a proportion of tracheid diameter), and stimulated formation of axial parenchyma cells in the xylem. Cryptomeria japonica seedlings adapted to flooding by forming adventitious roots, primarily on the original root system and submerged portion of the stem. Such new roots originated in the xylem ray parenchyma. Flooding stimulated ACC synthesis in roots and ethylene production in stems. The role of ethylene in alteration of stem anatomy is discussed.     

Temporal variation of microfibril angle in Eucalyptus nitens grown in different irrigation regimes

In 1990, a 2-ha plantation of Eucalyptus nitens (Deane and Maiden) Maiden was established in southeastern Tasmania and subjected to different irrigation regimes. Point dendrometers were installed in March 1995 to monitor radial stem movement every 15 min over several growing seasons. In this study, data from two growing seasons (1996-1998) were considered. From these measurements, daily increments of stem radius were determined. At the end of the second growing season, we extracted 12-mm cores and measured microfibril angles (MFA) of the wood at high resolution. Microfibril angles were rescaled on a time axis and mapped to daily and distance-based elements. Among treatments, irrigated trees in particular formed higher MFA early in the growing season (September-November) and lower MFA later in the growing season. Trees subjected to cyclic droughts showed clear relationships between MFA and soil water deficits, with MFA increasing in response to water stress release. Increases in MFA were preceded by accelerations in daily increment of stem radius. Among treatments, trees subjected to severe drought had the smallest MFA and generally low fluctuations in MFA. Irrigated trees were susceptible to changes in climate, whereas growth of the trees in the other treatments was limited by water availability. Use of path-analysis showed that temperature had an effect on stem radius increment but not on MFA; wind speed was the only factor that influenced MFA directly. Microfibril angle was correlated with stem shrinking and expansion phases; growth period length and growth rates were positively related to MFA.     

Radial stem growth in response to microclimate and soil moisture in a drought-prone mixed coniferous forest at an inner Alpine site

Dendroclimatological studies in a dry inner Alpine environment (750 m a.s.l.) revealed different growth response of co-occurring coniferous species to climate, which is assumed to be caused by a temporal shift in wood formation among species. The main focus of this study therefore was to monitor intra-annual dynamics of radial increment growth of mature deciduous and evergreen coniferous species (Pinus sylvestris, Larix decidua and Picea abies) during two consecutive years with contrasting climatic conditions. Radial stem growth was continuously followed by band dendrometers and modelled using Gompertz functions to determine time of maximum growth. Histological analyses of tree ring formation allowed determination of temporal dynamics of cambial activity and xylem cell development. Daily fluctuations in stem radius and radial stem increments were extracted from dendrometer traces, and correlations with environmental variables were performed. While a shift in temporal dynamics of radial growth onset and cessation was detected among co-occurring species, intra-annual radial growth peaked synchronously in late May 2011 and early June 2012. Moist atmospheric conditions, i.e. high relative air humidity, low vapour pressure deficit and low air temperature during the main growing period, favoured radial stem increment of all species. Soil water content and soil temperature were not significantly related to radial growth. Although a temporal shift in onset and cessation of wood formation was detected among species, synchronous culmination of radial growth indicates homogenous exogenous and/or endogenous control. The close coupling of radial growth to atmospheric conditions points to the importance of stem water status for intra-annual growth of drought-prone conifers.     

Effect of radial increment core diameter on tracheid length measurement in Norway spruce

The effect of radial increment core diameter (comparing the effects of 12, 8 and 4 mm diameters, with sampling from chips used as a control) on the measurement of tracheid mean length and the percentage of broken tracheids was examined for mature wood of Norway spruce. Two methods of measuring tracheid length were used: image analysis, in which arithmetic mean length (AML) was calculated for unbroken tracheids, and the Kajaani FS-200, in which AML was calculated for all tracheids, regardless of damage. In addition to AML, length weighted mean length (LWML) was calculated for the Kajaani. Almost 70 % of all tracheids remained undamaged when the 12 mm increment core was used. This was 18 % units less than for chips. Proportion of broken tracheids increased as core diameter decreased, and for the 4 mm increment core 30 per cent of all tracheids remained undamaged. All increment core diameters tested gave LWML values that were significantly lower than the control. For AML by Kajaani, all mean values were about 50 percent lower than corresponding LWML or image analysis values, thus showing the direct influence of broken tracheids. However, only the 4 and 8 mm increment cores differed significantly from the control. For unbroken tracheids measured by image analysis, the 4 mm increment core gave a significantly lower value than the control, thus showing the indirect influence of broken tracheids. Values obtained from the 8 and 12 :mm increment cores did not differ significantly from the control.     

How timing of stem girdling affects needle xylem structure in Scots pine

While needles represent a proportionally large fraction of whole-plant hydraulic resistance, no studies to date have investigated how source–sink disturbances affect needle xylem structure. In this study, we evaluated structural changes in xylem in current-year needles of Scots pine 227 and 411 days after stem girdling (hereafter referred to as DAG). Maximum and minimum tracheid lumen diameters and therefore also the size of tracheid lumen areas increased in needles 227 DAG compared to control needles. In contrast, tracheid dimensions were similar in needles 411 DAG as in the control needles, but smaller xylem area and lower number of tracheids resulted in the lower theoretical needle hydraulic conductivity of those needles. Several needle xylem parameters were intercorrelated in both control and girdled trees. These observed changes provide a new understanding of the processes that occur following a source–sink disturbance. Considering anatomical parameters such as the number of tracheids, tracheid dimension, or needle xylem area, which are rarely described in physiological studies, could be helpful, for example, in understanding to tree hydraulic systems or for modeling gas exchange. Finally, empirical equations were developed to calculate needle theoretical hydraulic conductivity and the number of tracheids in needles using an easily measurable parameter of needle xylem area.     

Weather factors controlling growth of Oriental beech are on the turn over the growing season

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.     

Comparing the intra-annual wood formation of three European species (Fagus sylvatica, Quercus petraea and Pinus sylvestris) as related to leaf phenology and non-structural carbohydrate dynamics

Monitoring cambial phenology and intra-annual growth dynamics is a useful approach for characterizing the tree growth response to climate change. However, there have been few reports concerning intra-annual wood formation in lowland temperate forests with high time resolution, especially for the comparison between deciduous and coniferous species. The main objective of this study was to determine how the timing, duration and rate of radial growth change between species as related to leaf phenology and the dynamics of non-structural carbohydrates (NSC) under the same climatic conditions. We studied two deciduous species, Fagus sylvatica L. and Quercus petraea (Matt.) Liebl., and an evergreen conifer, Pinus sylvestris L. During the 2009 growing season, we weekly monitored (i) the stem radial increment using dendrometers, (ii) the xylem growth using microcoring and (iii) the leaf phenology from direct observations of the tree crowns. The NSC content was also measured in the eight last rings of the stem cores in April, June and August 2009. The leaf phenology, NSC storage and intra-annual growth were clearly different between species, highlighting their contrasting carbon allocation. Beech growth began just after budburst, with a maximal growth rate when the leaves were mature and variations in the NSC content were low. Thus, beech radial growth seemed highly dependent on leaf photosynthesis. For oak, earlywood quickly developed before budburst, which probably led to the starch decrease quantified in the stem from April to June. For pine, growth began before the needles unfolding and the lack of NSC decrease during the growing season suggested that the substrates for radial growth were new assimilates of the needles from the previous year. Only for oak, the pattern determined from the intra-annual growth measured using microcoring differed from the pattern determined from dendrometer data. For all species, the ring width was significantly influenced by growth duration and not by growth rate, which differs from previous studies. The observed between-species difference at the intra-annual scale is key information for anticipating suitability of future species in temperate forests.     

Are neighboring trees in tune? Wood formation in Pinus pinaster

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.     

Diurnal and seasonal changes in stem increment and water use by yellow poplar trees in response to environmental stress

To evaluate indicators of whole-tree physiological responses to climate stress, we determined seasonal, daily and diurnal patterns of growth and water use in 10 yellow poplar (Liriodendron tulipifera L.) trees in a stand recently released from competition. Precise measurements of stem increment and sap flow made with automated electronic dendrometers and thermal dissipation probes, respectively, indicated close temporal linkages between water use and patterns of stem shrinkage and swelling during daily cycles of water depletion and recharge of extensible outer-stem tissues. These cycles also determined net daily basal area increment. Multivariate regression models based on a 123-day data series showed that daily diameter increments were related negatively to vapor pressure deficit (VPD), but positively to precipitation and temperature. The same model form with slight changes in coefficients yielded coefficients of determination of about 0.62 (0.57-0.66) across data subsets that included widely variable growth rates and VPDs. Model R2 was improved to 0.75 by using 3-day running mean daily growth data. Rapid recovery of stem diameter growth following short-term, diurnal reductions in VPD indicated that water stored in extensible stem tissues was part of a fast recharge system that limited hydration changes in the cambial zone during periods of water stress. There were substantial differences in the seasonal dynamics of growth among individual trees, and analyses indicated that faster-growing trees were more positively affected by precipitation, solar irradiance and temperature and more negatively affected by high VPD than slower-growing trees. There were no negative effects of ozone on daily growth rates in a year of low ozone concentrations.     

Temporal water deficit and wood formation in Cryptomeria japonica

Cell behavior in the cambium and developing xylem of 3-year-old Japanese cedar (Cryptomeria japonica D. Don.) trees, during and after an 11-day suspension of irrigation, was analyzed. Leaf xylem pressure potential and tangential strain of the stem surface were monitored throughout the experiment. Anatomical features and numbers of developing tracheids and cambial cells were observed in four trees, sampled on Days 0, 4, 8 and 11 after irrigation was suspended. Daytime xylem pressure potential decreased to -1.9 MPa on Day 7 and remained the same until irrigation was resumed on Day 11. The transverse dimensions of the tracheids, which began to form secondary walls, began to decrease on Day 4. The number of cells in the cambial zone and cell expansion zone decreased abruptly on Day 8. Tangentially aligned developing tracheids with collapsed cell walls were observed in samples harvested on Days 8 and 11. Secondary wall formation was recognized in these tracheids. After the resumption of irrigation, xylem pressure potential recovered rapidly to the same value as before the suspension of irrigation. Tangential strain increased within 30 min after the resumption of irrigation, and continued to increase until the onset of light the next day. Eighteen days after the resumption of irrigation, anatomical features of cells in the cambium and cell-expansion zone were similar to those observed before suspension of irrigation.     

Modelling day-to-day stem diameter variation and annual growth of balsam fir (Abies balsamea (L.) Mill.) from daily climate

In the present context of global climate changes and the continuous development of forest management strategies based on the concept of sustainable use, it is important to develop a better understanding of the climatic factors controlling the growth of boreal forests. In this study, we report the results of a five-year field research within which day-to-day balsam fir (Abies balsamea (L.) Mill.) stem diameter variation was measured with dendrometers and examined in relation with various daily climatic variables. A model built with data from three growing seasons that included solar radiation, relative humidity, temperature and precipitation explained 84% of the variance in day-to-day stem diameter variation from June to September. The model has approximately the same predictive capability when validated with independent daily data from two other growing seasons. The model captured both the cumulative increment associated with the irreversible growth and the high frequency variation of day-to-day fluctuations associated to changes in the stem water content. In general, rainy days during which relative humidity was high and solar radiation was low favored stem diameter expansion (growth and swelling) while stem diameter decreased during periods of low relative humidity and high solar radiation. Similar models were obtained when the June-September period was divided into two parts (June-July and August-September) to better represent the period during which most of the cumulative annual stem increment is observed (June-July). Inter-annual variation in stem growth computed from the modeled day-to-day variation in stem diameter was significantly correlated to the inter-annual variation in annual growth determined from tree core measurements over a 10 year period (p = 0.023). The model was notably able to capture a particularly poor growing year (2006) presumably due to a short-term heat stress period. Results suggest that the inclusion of daily data in growth-climate models may contribute to improve predictions of the potential tree growth response to climate by identifying particular climatic events that may escape to a classical dendroclimatic approach.     

Intra-annual variations in climate influence growth and wood density of Norway spruce

Intra-annual radial growth variations of two Norway spruce trees (Picea abies (L.) Karst.) were monitored over 4 years, at four heights up the stem, by means of point-dendrometers. The trees were then felled and radial wood samples were cut from the radii that had been monitored by the dendrometers and analyzed for density. From the radial growth measurements recorded by the dendrometers, we related positions within the rings to dates, thus making possible investigation of the relationships between changes within the rings in wood density and fluctuations in climate or growth rate. Radial growth started in early April and ended, with large intra-annual differences, in August or September. Short-term variations in growth rate were related to fluctuations in climate parameters and soil water reserves. The sensitivity of radial growth to climate decreased with stem height. Wood density responded strongly to drought events, and a dry period in June 1996 induced false-ring formation. Wood density was relatively independent of growth rate and climatic conditions during the first part of the growing season, but increased with decreasing radial growth rate later in the growing season.     

Effects of applied gibberellins and uniconazole-P on gravitropism and xylem formation in horizontally positionedFraxinus mandshurica seedlings

The present study deals with the effects of gibberellins (GA₃ GA₄) and uniconazole-P, an inhibitor of gibberellin biosynthesis, on negative gravitropism and xylem formation in the stems of horizontally positioned, 2-year-oldFraxinus mandshurica Rupr. var.japonica Maxim. seedlings. Each growth regulator (100 g) dissolved in 5 l acetone (50%) was applied to the basal node of the current shoot on May 24, 1995. The same treatment was repeated five times weekly until June 28. Five seedlings were used for each treatment. The seedlings were positioned horizontally 24h after the first application on May 25. Within 5 weeks the horizontal stem of control and GA-treated seedlings exhibited negative gravitropism. In contrast, the application of uniconazole-P inhibited negative gravitropic stem bending. The application of GAs increased the number of gelatinous fibers having thickened cell walls on the upper side of stems. The uniconazole-P application decreased xylem cell formation but did not inhibit the formation of gelatinous fibers. These results indicate that not only the differentiation of gelatinous fibers but also xylem increment is important in the negative gravitropism of horizontally positionedF. mandshurica seedlings. These results also suggest that GAs may be involved in xylem cell :formation rather than the differentiation of gelatinous fibers in this species.Part of this research was reported at the 23rd annual meeting of the Plant Growth Regulation Society of America, Calgary, Canada, July 1996     

Lignin deposition during earlywood and latewood formation in Scots pine stems

Lignin deposition at consecutive secondary wall thickening stages of early and late xylem cells during annual ring wood formation in Scots pine (Pinus sylvestris L.) stems was studied. Lignin patterns, isolated by thioglycolic acid method, consisted of alcohol-soluble (LTGA-I) and alkali-soluble (LTGA-II) fractions. The sum of two fractions, being the total lignin content, gradually increased in the course of lignification. However, the increments of lignin amount at each development stage of early and late tracheids were different. The intensity of lignin deposition increased in the course of earlywood tracheid maturation and decreased toward the end of latewood cell differentiation. The deposition of two lignin fractions in each layer of forming wood also occurred oppositely. The increment of LTGA-I descended, whereas that of LTGA-II increased from the beginning to the end of early xylem lignification. In contrast, LTGA-I increment dropped, whereas LTGA-II rose during late xylem lignification. Gel permeation chromatography showed that the lignins, formed at the beginning of lignification, were more homogeneous and had higher molecular weight compared with the lignins at the end of cell differentiation. Besides, the content of cellulose, estimated as the residue after lignin isolation, and of cell wall substances, presented as cell wall cross-section areas, at consecutive maturation stages of early and late xylem cells have been found to be different. The data show that lignin deposition occurred in different conditions and with opposite dynamics during early and late xylem formation.     

Effects of xylem cavitation and freezing injury on dieback of yellow birch (Betula alleghaniensis) in relation to a simulated winter thaw

Shoot dieback, shoot growth, stem xylem cavitation, stem and root freezing injury, and root pressure were measured in 2-year-old, cold-hardened, potted yellow birch (Betula alleghaniensis Britt.) seedlings that had been subjected to a simulated winter thaw for 0, 5, 10, 19 or 27 days followed by 10 weeks at -10 degrees C. Stem xylem cavitation was determined as percent loss of hydraulic conductivity. Stem freezing injury was measured as electrolyte leakage (EL). Root freezing injury was determined by EL and by triphenyl tetrazolium chloride (TTC) reduction. Thaw duration was significantly correlated with dieback, new shoot growth, stem xylem cavitation, stem and root freezing damage, and root pressure (P < 0.05). In particular, shoot dieback was positively correlated with stem xylem cavitation (P < 0.001), residual stem xylem cavitation (P < 0.01) and root freezing injury (P < 0.010), but only weakly correlated with stem freezing damage (P < 0.05). In roots, freezing damage was negatively correlated with root pressure (P < 0.05), which, in turn, was negatively correlated with residual stem xylem cavitation after root pressure development. In stems, there was no correlation between freezing damage and xylem cavitation. We conclude that long periods of winter thaw followed by freezing resulted in freezing injury to roots concomitant with a reduction in root pressures, leading to poor recovery from freezing-induced xylem embolism.     

银中杨生长动态分析及模型建立

通过对银中杨林木生长模型的建立 ,确定出银中杨树高生长速增期为 7月 6日至 2 8日 ,速增点为 7月 17日 ;胸径速增期为 6月 2 1日至 8月 2 9日 ,速增点为 7月 2 5日 ;材积生长速增期为 11～ 17a,速增点为第 14年 ,从而揭示了银中杨林木生长随时间的动态变化规律