毛白杨形成层的活动周期及其POD同工酶的变化

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

Photosynthate distribution patterns in cherrybark oak seedling sprouts

We used 14C tracers to determine photosynthate distribution in cherrybark oak (Quercus pagoda Raf.) seedling sprouts following release from competing mid-story vegetation. Fall acquisition of labeled photosynthates by seedlings followed expected source-sink patterns, with root and basal stem tissues serving as the primary sinks. Four months after the seedlings had been labeled with 14C, they were clipped to induce sprouting. First-flush stem and leaf tissues of the resulting seedling sprouts were the primary sinks for labeled photosynthates stored in root tissues. Second-flush stem and leaf tissues, and first-flush stem and leaf tissues the following growing season, were not primary sinks for labeled photosynthates stored in root tissues despite the high radioactivity in root tissues. Root tissues appeared to deposit photosynthates in a layering process whereby the last photosynthates stored in new xylem were the first to be depleted during the initiation of a growth flush the following spring. There were more labeled photosynthates in roots of released seedling sprouts compared with non-released seedling sprouts, indicating increased vigor of released seedling sprouts in response to greater light availability. In contrast, stem and source leaf tissues of non-released seedling sprouts contained greater percentages of labeled photosynthates compared with released seedling sprouts, indicating either greater sink strength or poorly developed xylem and phloem pathways that created inefficiencies in distribution to root tissues. The 14C distribution coefficients confirmed the distribution patterns and provided additional information on the important sinks in released and non-released cherrybark oak seedling sprouts.     

Frost tolerance and bud dormancy of container-grown yellow birch, red oak and sugar maple seedlings

Container-grown seedlings of red oak (Quercus rubra L.), sugar maple (Acer saccharum Marsh.) and yellow birch (Betula alleghaniensis Britton) in their first year of growth were overwintered outdoors. Tolerance of roots and stems to freezing was compared from late summer to the following spring. Mitotic activity in the apical bud was related more closely to air temperature than to bud dormancy as defined by days to bud break. In all species, stem hardening was observed before days to bud break reached a maximum. Dormancy release (days to bud break equal to zero) of yellow birch coincided with loss of stem hardening in the spring. Roots hardened more slowly, had a lower frost tolerance than stems in fall and winter, and dehardened earlier than stems in the spring. There were differences in stem and root hardiness among the species, with yellow birch being the most tolerant, followed by sugar maple and red oak. Primarily because of root sensitivity to frost, winter was a critical period for all three species, but particularly for red oak.     

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.     

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.     

Cellular structure and chemical composition of cork from the Chinese cork oak (Quercus variabilis)

Quercus variabilis Blume, the Chinese cork oak, is an oak species with a thick cork outer bark. The cork is exploited at a limited scale in China and considered of lower quality than the commercial cork from Quercus suber. We studied an industrial cork granulate feedstock of Q. variabilis in relation to cellular structure and chemical composition and compared it to Q. suber cork under a material’s perspective. The cork of Q.variabilis has 1.1 % ash, 9.6 % extractives, 34.8 % suberin and 19.1 % lignin. The monosaccharide composition with shows a predominance of hemicelluloses: glucose 42.8 % of total neutral sugars, xylose 27.5 %, arabinose 15.4 %, galactose 9.0 %, mannose 4.0 %, rhamnose 1.2 %. The FT-IR spectrum shows the indicative peaks of suberin. The composition is overall similar to that of Q. suber cork. Q. variabilis cork has the typical cellular characteristics of bark cork tissues with a regular and radially aligned structure of cells without intercellular voids. Solid volume fraction was estimated at approximately 16 %. Compared with Q. suber, the Q. variabilis cork cells are smaller, the cell wall undulation and the overall row alignment less homogeneous, the cell surface is irregular and the solid volume proportion higher. The characteristics of Q. variabilis cork support its use as a cellular material for sealing, insulation and energy absorption, but the overall quality is lower than that of Q. suber cork. The negative impact of the higher density and structural lower uniformity at tissue and cell level should be evaluated for processing and product performance.     

Opportunity for hybridization between two oak species in mixed stands as monitored by the timing and intensity of pollen production

The opportunity of cross-pollination in mixed stands of two oak species (cork oak and holm oak) was studied by characterizing individual phenologies of flowering. In the spring of 1998 at one stand consisting of 64 marked trees, there was a period of 19 days when maximal pollen release in one species and stigma receptivity in the other occurred simultaneously, enabling interspecific gene flow in either direction. This happened in spite of an average time separation of 22 days between the two species, reflecting a considerable intraspecific variation in the timings of flowering. Flowering intensities (as estimated from male flower abundance) were high, but fruiting intensities were comparatively low. Shortly after pollination, considerable abortion of female flowers and early fruits was recorded. In 2000, the interspecific overlap of phenologies was drastically reduced due to a delay in cork oak flowering. On the other hand, the individual timings were repeatable for most trees, at least in holm oak. Two other mixed stands were subject of parallel studies, with similar results in all traits except for a less dramatic reduction in fruiting intensities. In spite of the high opportunity for cross-pollination in 1998, and given the lack of hybrids among the progenies from the subsequent fruiting season [Oliveira, P., Custódio, A.C., Branco, C., Reforço, I., Rodrigues, F., Varela, M.C., Meierrose, C., 2003. Hybrids between cork oak and holm oak: isoenzyme analysis. Forest Genet. 10, 283–298], it can be concluded that the prerequisite of cross-pollination is clearly insufficient for hybridization to succeed. Post-pollination processes must play an important role in the maintenance of reproductive isolation between the two species.     

Physiological responses of cork oak and holm oak to infection by fungal pathogens involved in oak decline

The aim of this research was to study the changes in net photosynthesis and stomatal conductance values in 3‐year‐old cork oak and holm oak seedlings growing in natural conditions and inoculated with Apiognomonia quercina, Biscogniauxia mediterranea, Botryosphaeria corticola and Pleurophoma cava. Throughout the 4‐month experimental period, the evolution of visual external symptoms and the values of physiological variables were periodically recorded. All pathogens caused stem lesions around the infection point; however, the lesions caused by B. corticola were longer in both oak species. On cork oak seedlings, all pathogens induced a significant and gradual reduction in net photosynthesis and stomatal conductance values, whereas other physiological disturbances were induced only by B. corticola infections on holm oak seedlings.     

Effects of partial defoliation on carbon and nitrogen partitioning and photosynthetic carbon uptake by two-year-old cork oak (Quercus suber) saplings

At the end of the growing season in late July, 20-month-old cork oak (Quercus suber L.) saplings were partially defoliated (63% of leaf area) to evaluate their ability to recover leaf area after defoliation. At 18 and 127 days after defoliation, changes in starch and nitrogen pools were determined in leaves and perennial organs, and variations in photosynthetic carbon uptake were investigated. To determine the role of stored nitrogen in regrowth after defoliation, plant nitrogen was labeled in the previous winter by enriching the nutrient solution with 15N. Plants recovered the lost leaf area in 127 days. Although there was remobilization of starch and nitrogen from leaves and perennial organs, the availability of resources for growth in the following spring was not decreased by defoliation. On the contrary, starch concentration in coarse roots was higher in defoliated saplings than in control saplings, presumably as a result of the higher net CO2 exchange rate in newly developed leaves compared with pre-existing leaves.     

The effect of cork removal on the radial growth and phenology of young cork oak trees

The effect of the removal of cork was studied in 11-years old cork oak trees (Quercus suber L.), growing in favorable conditions, in relation to phenology and radial growth during two years. Longevity of leaves was 14–15 months (1996, 1998) and 10 months (1997). Bud burst started in mid-February and leaf flushing in April, extending until June. Neither a distinct two-period flushing nor an autumn bud burst or leaf flushing were observed. Radial growth started in mid-April and continued until the end of November, with the maximum growth in June and July. In trees with the removal of cork, leaf abscission occurred a little earlier and new branches had on average 23% fewer new leaves. The radial growth of the trees and its general seasonal pattern were not affected by cork removal in the year of removal or in the year after. The only observation was a shift of the maximum radial growth rate from June to July for the trees where cork had been removed.     

Peridermium pini (Pers.) Lev. and the Resin-top Disease of Scots Pine: II. Lesion Anatomy

The stages in the formation of Peridermium pini lesions aredescribed. The hyphae of the parasite were septate, uninucleate,and intercellular, with unbranched haustoria penetrating allphloem cells and xylem parenchyma cells. Lesion extension occurredby apical growth of the hyphae along the outer edge of the functionalphloem. The hyphae stretched unbroken across the active cambium.Active centripetal and longitudinal growth of the mycelium occurredin the resin ducts of the xylem, the hyphae commonly reachingthe pith. Spermogonia were produced in late summer and fall,aecia appeared in spring. In lesion xylem, tracheids were shorter,rays taller, more numerous and wider, but ray cell height remainedunchanged. Fusiform rays increased in number and size and verticalresin ducts increased in number. Changes in xylem tissues didnot become evident until two to four years after invasion ofthe cambium by the mycelium. Invasion of the secondary phloemresulted in an increase in number and size of the longitudinalphloem parenchmya cells, while changes in phloem rays were similarto those in xylem rays.     

Thermal optima of photosynthetic functions and thermostability of photochemistry in cork oak seedlings

Temperature effects on photosynthesis were studied in seedlings of evergreen Mediterranean cork oak (Quercus suber L.). Responses to changes in temperature and the temperature optima of maximal carboxylation rate (V(cmax)) and maximal light-driven electron flux (J(max)) were estimated from gas exchange measurements and a leaf-level photosynthesis model. The estimated temperature optima were approximately 34 and 33 degrees C for V(cmax) and J(max), respectively, which fall within the lower range of temperature optima previously observed in deciduous tree species. The thermostability of the photosynthetic apparatus was estimated according to the temperature at which basal chlorophyll a fluorescence begins to increase (T(c)). The T(c) was highly variable, increasing from 42 to 51 degrees C when ambient temperature rose from 10 to 40 degrees C, and increasing from 44 to 54 degrees C with decreasing soil water availability while net CO(2) assimilation rate dropped to almost zero. When a heat shock was imposed, an additional small increase in T(c) was observed in drought-stressed and control seedlings. Maximal T(c) values following heat shock were about 56 degrees C, which, to our knowledge, are the highest values that have been observed in tree species. In conclusion, the intrinsic temperature responses of cork oak did not differ from those of other species (similar T(c) under ambient temperature and water availability, and relatively low thermal optima for photosynthetic capacity in seedlings grown at cool temperatures). However, the large ability of cork oak to acclimate to drought and elevated temperature may be an important factor in the tolerance of this evergreen Mediterranean species to summer drought and high temperatures.     

Oak restoration in water-limited pine plantations: interactive effects of overstory light interception and water availability on understory oak performance

Oak regeneration within pine monocultures is an opportunity to diversify forest structure. We examined the relationships between overstory (Pinus brutia) light interception and understory oak (Quercus ithaburensis) performance in water-limited forests. The study was performed in a mature pine plantation in Mediterranean Israel. Twenty-year-old oaks differing in location with respect to pine overstory and representing a gradient of light availability, such as open space (irradiance 100 %), interface (17–77 %), and understory (14–23 %), were monitored. Photosynthetic photon flux density (PPFD), leaf gas exchange, and twig water potential (TWP) were measured during the growth season under increasing drought stress. Predawn TWP decreased sharply from early to late spring and was positively related to irradiance during mid-spring only. Predawn to midday TWP gradient was positively related to irradiance mostly so during mid-spring. Daily averages of stomatal conductance (gs), net carbon assimilation rate (A), and transpiration rate (E) were highest in early spring and decreased gradually toward late spring. They were positively related to irradiance though this relationship became less pronounced from early to late spring. Oak height and stem basal area were positively related to irradiance. A/gs ratio was positively related to irradiance throughout the entire growth season. It increased from early to mid-spring but decreased toward late spring. A/PPFD ratio decreased from early to late spring showing a negative relationship with irradiance. We concluded that light availability was mainly responsible for spatial variation in oak performance and proposed that small-scale overstory gaps aiming for direct sunlight exposure during early spring should achieve maximum understory oak performance with minimal pine removal.     

Contrasting distribution and seasonal dynamics of carbohydrate reserves in stem wood of adult ring-porous sessile oak and diffuse-porous beech trees

We tested the hypothesis that broad-leaved forest species with contrasting wood anatomy and hydraulic system (ring-porous versus diffuse-porous) also differ in distribution and seasonal dynamics of carbohydrate reserves in stem wood. Total nonstructural carbohydrate (TNC) reserves (starch and sugars) were measured enzymatically in the 10 youngest stem xylem rings of adult oak (Quercus petraea (Matt.) Liebl.) and beech (Fagus sylvatica L.) trees during an annual cycle. Radial distribution of carbohydrates was investigated according to ring age. On all dates, oak trees had twofold higher TNC concentration than beech trees (41 versus 23 mg g(DM)(-1)), with starch accounting for the high TNC concentration in oak. Seasonal dynamics of TNC concentration were significantly (P < 0.05) more pronounced in oak (20-64 mg TNC g(DM)(-1)) than in beech (17-34 mg TNC g(DM)(-1)). A marked decrease in TNC concentration was observed in oak trees during bud burst and early wood growth, whereas seasonal fluctuations in TNC concentrations in beech trees were small. The radial distribution of TNC based on ring age differed between species: TNC was restricted to the sapwood rings in oak, whereas in beech, it was distributed throughout the wood from the outermost sapwood ring to the pith. Although the high TNC concentrations in the outermost rings accounted for most of the observed seasonal pattern, all of the 10 youngest xylem rings analyzed participated in the seasonal dynamics of TNC in beech trees. The innermost sapwood rings of oak trees had low TNC concentrations. Stem growth and accumulation of carbon reserves occurred concomitantly during the first part of the season, when there was no soil water deficit. When soil water content was depleted, stem growth ceased in both species, whereas TNC accumulation was negligibly affected and continued until leaf fall. The contrasting dynamics and distribution of carbohydrate reserves in oak and beech are discussed with reference to differences in phenology, early spring growth and hydraulic properties between ring-porous trees and diffuse-porous trees.     

Cambial Anatomy and Annual Rhythm of Secondary Xylem Development in the Twigs of Azadirachta indica A. Juss. (Meliaceae) Growing in Different Forests of Gujarat State

Abstract

Seasonal behavior of vascular cambium and development of secondary xylem were studied in the 2-3-years-old branches of Azadirachta indica A. Juss. (Meliaceae) growing in moist deciduous (MDF), dry deciduous (DDF) and scrub land (SF) forests of Gujarat State. Development of xylem began with the sprouting of new leaves in January in both MDF and DDF. Xylem production culminated in July in MDF whereas it reached peak in April and July in DDF. Cambial cells ceased to divide with the maturation of leaves in November and October in MDF and DDF, respectively. Mature xylem and phloem derivatives surrounded the cambium in December in MDF and November in DDF. Radial growth in the branches of trees growing in SF was found continuous throughout the year with peak cambial activity in April, July and November. In all the three forests, maximal radial growth was encountered in July when the rains were heavy. Cambial activity and xylem development showed significant correlation with the phenology of the trees in MDF and DDF whereas cambium was found active throughout the year and no correlation was found between the cambial activity and phenology in SF.     

Natural decline of volatile acids due to aging of living trees

Wood age, i.e., number of years that have passed after the annual rings of xylem tissue developed from cambium, had a strong negative impact on the volatile acid content of wood of the two oak species, Quercus petraea and Q. frainetto. Up to 80% of acetyl groups were cleaved off in the heartwood zones close to the pith if the living oak trees were about 200 years old.     

Development of cork oak (Quercus suber L.) seedlings in response to tree shelters and mulching in northwestern Tunisia

The need for reforestation in cork oak (Quercus suber L.) areas is challenged by difficulties. Principal among these is herbivory of young plants, vegetative competition, and slow growth rates of cork oak seedlings. We evaluated the early development of cork oak seedlings treated using tree shelters and mulching in northwestern Tunisia. We tested three tree shelter treatments (non-vented, vented, and control) to shield seedlings from animal damage and five mulch types to control competing vegetation (Italian Stone Pine, Lentisk, combination of Italian Stone Pine and Lentisk (organic mulches), gravel (inorganic mulch) and no mulch). At the end of the two-year experiment, sheltered seedlings were 89-99% taller than unshelteredseedlings and had higher numbers and lengths of shoot growth flushes. In contrast, both stem diameter growth and dry weight biomass (from samples extracted after two years) were significantly reduced inside tree shelters. Root-to-shoot ratio was not significantly different in sheltered vs. unsheltered seedlings, suggesting that tree shelters do not adversely affect this parameter. Mulching alone did not favour growth, but could be beneficial when combined with tree shelters. The combination of vented tree shelters and gravel mulch was the most effective treatment for promoting diameter, height and stem volume growth.     

Regular cambial activity and xylem and phloem formation in locally heated and cooled stem portions of Norway spruce

The effect of heating (23–25°C) and cooling (9–11°C) on regular cambial activity and xylem and phloem formation in the stem portion of Norway spruce was investigated. Adult trees were sampled at 21-day intervals during the 2005 vegetation period. Continuously elevated temperatures increased the rate of cell division in the first part of the growing season, but did not significantly prolong cambial activity at the end of the vegetation period in the heated tree. Low-temperature treatments shortened regular cambial activity and slowed down cell production. The xylem growth ring was wider in the heated sample and narrower in the cooled sample compared to the control. The temperature in the cambial region was only negligibly transferred along the stem from the site of its application. In general, the temperature in the cambium was affected by a long-term rise or drop in air temperatures. Both experiments affected the structure and width of phloem growth increments. The tangential band of the axial parenchyma was not continuous in the cooled sample. The number of late phloem cells was reduced in the cool-treated sample and increased in the heat-treated sample. Our experiments confirmed the effect of constantly increased or decreased temperatures on regular cambial activity in Norway spruce.     

Cork oak (<Emphasis Type="Italic">Quercus suber</Emphasis> L.) wood growth and vessel characteristics variations in relation to climate and cork harvesting

Variations in tree ring growth of Quercus suber L. were analysed using dendrochronological techniques on cork oak discs from trees harvested in the cork producing region of Alentejo, Portugal. A tree-ring chronology containing a strong common signal and covering the period from 1970 to 1995 was build for ca. 30-year-old cork oaks never submitted to cork harvesting using 14 trees that crossdated satisfactorily out of 30 sampled trees. The tree ring indices correlated positively with September temperature (r = 0.48, P < 0.05) and very strongly with precipitation totals from previous October until current February (r = 0.82, P < 0.001) showing that the water stored in the soil during the autumn and winter months prior to the growing season has a primordial effect on the growth of the given season. The effects of cork harvesting were analysed by comparing mean ring width, mean annual vessel area, vessel density (n°vessels/mm²), and vessel coverage (percentage of transverse surface occupied by vessels) between three mature cork oak trees and three young trees, for the period from 1987 to 1996, corresponding to the growth between two consecutive cork removals in the case of mature trees. In 1988, 1989 and 1996 (corresponding to the first and second years after cork removal, and 1996 to a year of cork removal), the ratios between ring widths of young versus mature trees was twice that for the rest of the period. However, an effect of cork removal indicated by eventual alterations in vessel size and distribution in the wood rings corresponding to the years 1988, 1989 and 1996 in the mature cork oaks was not observed.