云南轿子山不同海拔急尖长苞冷杉径向生长动态及其低温阈值

环境对树木的功能和生存具有至关重要的意义, 但环境因素影响树木发育的过程仍有待探究。树木径向生长动态监测是深刻了解树木生长状况及其对气候变化响应的重要手段。该研究以云南轿子山不同海拔树线树种急尖长苞冷杉(Abies georgei var. smithii)为研究对象, 对其年内径向生长动态进行监测, 以期明晰树木径向生长各阶段起止时间的海拔差异, 分析不同海拔树木形成层活动和木质部分化对温度的响应, 辨析树木径向生长的低温阈值。结果表明: (1)海拔越高, 径向生长开始的时间越晚, 结束越早, 生长季缩短。海拔每上升100 m, 急尖长苞冷杉径向生长的开始时间推迟4.7 d, 结束时间提前7.2 d, 生长季缩短12.8 d; (2) 3个海拔急尖长苞冷杉的径向生长具有相似的低温阈值(约5 ℃), 温度控制形成层活动的开始与木质部分化的结束; (3)不同海拔急尖长苞冷杉形成层的细胞分裂活动均在温度较高时(夏至日前后)减弱。 形成层活动的结束与海拔引起的温度变化关系较弱, 光周期可能参与了形成层活动结束的调控, 以确保当年新生细胞能够在冻害来临前完成木质化。该研究结果有助于加深树木生长动态对气候变化响应机制的认识, 为更好地适应和应对气候变化提供科学依据。     

树轮与交叉定年

树木的生长主要包括径向的生长（长宽）和纵向的生长（长高）。树木的生长除受内部因素的控制外,也受外部环境条件的制约。在具有显著季节变化的温带和寒带地区,树木一般每年都存在周期性的生长变化。春天,紧挨着木质部外面的形成层细胞开始分裂,向内形成木质部细胞,分裂后的细胞大而壁厚,颜色鲜嫩,被称为早材.也叫春材;以后细胞生长减慢,壁更厚,     

贺兰山地区植被冠层物候与树干形成层物候的关系

基于贺兰山地区98棵油松树轮样本的宽度数据、植被归一化指数(NDVI)数据以及土地覆被数据,采用VS-oscilloscope模型模拟的油松径向生长过程,研究植被冠层与树干形成层物候之间的联系。结果表明: 林地冠层与油松形成层生长结束期(EOS)显著相关,且高于草地与形成层之间的相关。油松生长开始期(SOS)和EOS分别与5—6月、8—9月的平均最低温度有关。5—6月的平均最低气温每升高1 ℃,SOS提前4.3 d;8—9月的平均最低气温每升高1 ℃,EOS推迟2.6 d。植被冠层物候与油松形成层物候的相关性受植被类型的影响;仅通过树轮生理模型模拟树木生长动态,结果可能存在偏差;利用遥感监测数据将冠层发育和形成层生长过程结合有助于更准确地了解树木生长动态。     

热带喀斯特森林多花白头树木质部年内生长动态及其对环境因子的响应

多花白头树(Garuga floribunda var.gamblei)是西双版纳热带喀斯特森林中常见的落叶树种，容易受水分亏缺的影响。为探究热带喀斯特森林树木年内生长动态及其对环境因子的响应，该研究利用生长环和微树芯两种方法监测了多花白头树在2020—2021年的树干径向变化和木质部生长动态，并分析了树干径向变化和木质部生长与环境因子的相关性。微树芯法的监测表明，在2020年，多花白头树在3月底开始出现扩大细胞，9月底细胞壁加厚结束；在2021年，多花白头树在4月中旬开始出现扩大细胞，10月中旬细胞壁加厚结束。生长环的监测表明，2020年和2021年的径向生长开始时间均晚于扩大细胞出现时间，径向生长结束时间均早于细胞壁加厚结束时间。多花白头树在2020年和2021年的木质部生长持续时间大致相同，2020年的木质部生长量((2.87±1.46)mm)也与2021年((2.98±1.02)mm)几乎一致。多花白头树的月径向生长量、扩大细胞区域宽度和细胞壁加厚区域宽度均与降水呈显著正相关，这表明水分状况在多花白头树的木质部生长过程中发挥了重要作用；扩大细胞区域宽度还与日平均气温和日最低温显...     

热带季节雨林多花白头树年内径向生长动态及其对环境因子的响应

为了解热带地区树木的季节性生长动态和规律,在西双版纳热带季节雨林利用高精度生长仪和微树芯法对落叶树种多花白头树的径向生长季节动态进行监测。结合木质部非结构性碳水化合物和环境因子的监测,分析其形成层活动和径向季节动态的生理生态驱动因子。结果表明: 在2020年,生长仪的监测显示,多花白头树于5月底(儒略日DOY:149.3±7.2)开始生长,8月底(DOY:241.0±14.7)生长结束,年生长量为3.12 mm,最大生长速率为0.04 mm·d^-1。而微树芯法显示,扩大细胞3月9日(DOY:69.2±6.2)开始出现,9月19日(DOY:262.8±2.8)细胞加厚结束,木质部生长量为1.76 mm,最大生长速率为0.009 mm·d^-1。多花白头树径向日生长量与生长季的降水、相对湿度、日最低气温、深度为20 cm的土壤含水量和温度呈显著正相关,而与日最高气温、水汽压亏缺、最大风速和水汽压呈显著负相关。多花白头树边材淀粉含量和可溶性糖含量均在生长季开始之前保持较高水平,淀粉含量在3月底达到最低,而可溶性糖含量5月中旬达到最低,随着生长季的结束淀粉和可溶性糖的含量分别在10月中旬和12月底达到最高。     

白皮松形成层的活动周期及其多糖贮量和淀粉酶同工酶的变化

4月初,白皮松(Pinus bungeana Zucc.)形成层带细胞开始增大,未成熟的木质部和韧皮部细胞增多,下旬出现成熟的木质部细胞。5月以后,木质部和韧皮部的形成速度加快,6月初进入晚材形成期。8月初停止产生木质部,9月中旬停止产生韧皮部。多糖颗粒的消长与形成层活动有较强的相关性,恢复活动前后颗粒含量持续增长,6月进入晚材形成期才持续减少,至翌年1月初完全消失,3月又重新积累,并迅速达到高峰。淀粉酶同工酶在活动期只有一条酶带,形成层停止产生木质部后出现了3条特异酶带,12月初又出现了2条特异酶带,这5条酶带都一直存在到形成层恢复活动。     

年轮和生长轮

木本植物茎的增粗是形成层细胞活动的结果。在四季分明的温带地区,由于季节变化对形成层的影响,通常每年形成一轮年轮。春天气候温和,雨水充沛、植物体生长代谢旺盛,适宜维管形成层活动,细胞的分裂和生长都快,所形成的木质部导管数目较多,腔大壁薄,而木纤维较少,因而这部分木质部质地疏松,颜色较浅,称为早材或春材。秋季,气温渐低,雨水渐少,植物的生长代谢变慢,形成层的活动减弱,生长的情形刚好相反,形成质地坚实致密,颜色较深的晚材,亦称夏材或秋材。裸子     

祁连山中部祁连圆柏年内径向生长对气候因子的响应

通过对祁连山中部葫芦沟流域的祁连圆柏连续采集微树芯,对其形成层活动和径向生长动态进行了连续两年的监测研究。结果表明,2012年细胞壁加厚和细胞成熟阶段开始时间分别发生在6月26日和7月24日,比2013年细胞壁加厚（6月22日）和细胞成熟阶段（6月26日）开始时间分别晚5 d和28 d。2012年细胞扩大、细胞壁加厚和细胞成熟阶段结束时间分别为7月16日、8月9日和9月8日,比2013年各阶段结束时间分别晚7、28 d和24 d。2012年最大细胞分裂速率为0.33细胞/d,共形成20.9个细胞,细胞分裂速率和木质部细胞总数均高于2013年。通过与附近气象站记录的气象数据进行对比,发现祁连圆柏生长开始时间在温暖年份显著早于寒冷年份,说明祁连圆柏的径向生长开始时间与温度有关。但2013年春季和夏初的高温导致区域干旱程度加剧,使祁连圆柏生长结束时间显著早于2012年,并导致2013年的木质部细胞总量和生长速率都小于2012年。研究表明,在寒冷干旱地区,尽管升温会使生长季提前,但升温导致的干旱胁迫可能对树木的生长速率和木质部细胞总量产生重要影响。     

树木木质部生长动态及其调节机制研究进展

全球变化对树木木质部生长产生了深远影响,进而影响了森林生态系统的固碳功能以及全球生态系统能量和物质的循环过程。树木木质部生长动态主要包括形成层活动开始和结束的时间、生长季长度以及分裂速率等,其受到多种因素的共同调节,如植物激素、碳水化合物、氮素和气象因子等。通过在精细的时间尺度上对比研究树木木质部生长动态,揭示木质部形成的决定因子,可以加深对树木生长生理机制的理解,从而提高其对气候变化响应的预测精度。对近年来在木质部的形成动态及其调节机制方面取得的研究进展进行了综述,并对未来的研究方向进行了展望。     

毛白杨维管形成层带细胞超微结构的季节性变化

木材(次生木质部)是树木形成层细胞分化的产物,形成层的活动方式不仅影响木材的产量,而且影响木材的结构和性质.利用透射电子显微镜观察了生长在北京地区的毛白杨(Populus tomentosa Carr.)枝条形成层带细胞一个完整活动周期的超微结构变化.在木质部母细胞完全恢复活动之前,形成层纺锤状原始细胞的分裂和韧皮部细胞的分化已经开始.枝条上芽的展开和幼叶的生长可能决定了形成层带细胞的这种活动方式.透射电镜观察更清楚地揭示了树木形成层细胞在活动初期的分化特点.活动期形成层细胞中的大液泡在进入休眠期后逐渐分成许多小液泡分散在细胞质中.随着液泡融合逐渐消失的深色蛋白类物质又重新充满了大部分液泡.油滴和淀粉颗粒的年变化情况同液泡中的蛋白类物质基本相似.无论在活动期还是休眠期,形成层纺锤形细胞的质膜上都发现有许多可能与物质运输有关的小泡状内折.由核膜、内质网和高尔基体及其分泌小泡组成的细胞内膜系统,在形成层活动周期的不同阶段,其形态和分布明显不同,尤其在形成层细胞的恢复活动及其衍生木质部细胞次生壁的沉积过程中发挥着重要作用.整个活动周期中,形成层纺锤形细胞的径向壁都比弦向壁厚,处在休眠期的形成层带细胞,其径向壁与弦向壁的差别则更明显.形成层恢复活动时,径向壁上特别是与弦向壁相连的角隅处出现部分自溶现象.细胞壁特别是径向壁的变薄是形成层细胞恢复活动的重要特征.     

暖干化加剧东北半干旱地区油松人工林径向生长的水分限制

相较天然林,人工林生态系统对全球性气候变化更敏感。本文利用树木年代学方法,以东北半干旱地区油松人工林为对象建立油松年轮宽度年表,研究油松生长的动态变化及其径向生长与气象因子的相关关系,探讨升温对油松生长及分布的影响。结果表明: 研究区油松年轮宽度主要与生长季5—7月的平均温度呈显著负相关,与生长季早期4月和生长季5—7月的平均降水量和PDSI呈显著正相关,水分可利用性是限制研究区油松径向生长的主要因子。自西南向东北随着年降水量增加,各样点油松径向生长对年均温的敏感性增强,与年降水量的相关关系由显著正相关转变为负相关,说明偏干旱的西南部地区油松生长受水分限制更严重。气候变暖导致的干旱胁迫使得研究区西南部的部分人工林油松生长呈衰退状态。随着暖干化的持续,研究区油松分布边界将发生局地收缩,适宜生长的边界将向北移动。     

How needle phenology indicates the changes of xylem cell formation during drought stress in Pinus sylvestris L.

Temperature, water availability and photoperiod are the primary drivers of tree phenological processes. However, there is lack of information how the spring temperature and summer drought affect phenology of both cambium and needles. We evaluated the interplay between xylem and needle development of mature Scots pine trees in three consecutive years differing in the spring temperatures and onset and duration of summer drought. Cambial activity began on the day of a year (DOY) 83–87, while the bud break occurred on DOY 113–119, when also the secondary cell wall of tracheids started its formation. While the timing of bud break correlated with the sum of effective temperatures, the beginning of cambial zone activity did not. The needles were fully unfolded around DOY 170, which corresponded to the transition between earlywood and latewood tracheids. Summer drought did not affect needle development, but it changed the rate of production and morphology of latewood tracheids. Latewood tracheids from the year 2015 when the drought was longest (i.e. until the second half of August in 2015 compared to the duration from June until the middle of July in the two other years) were 32% narrower and they had 34% thinner cell walls than in 2014 and 2016. The improvement of tree water status in July resulted in the formation of intra-annual density fluctuations (IADF). The interplay between needle and cambium phenology provided an insight into how the tree allocates the resources with varying temperature and soil water availability.     

Growth Rings,Phenology, Hurricane Disturbance and Climate in Cyrilla racemiflora L., a Rain Forest Tree of the Luquillo Mountains,Puerto Rico1

The growth phenology of Cyrilla racemiflora L., the dominant tree species of the montane rain forest, (subtropical lower montane rain forest, sensu Holdridge) of the Luquillo Mountains of Puerto Rico was studied intensively during 1989, and then semiannually through mid-1993 to determine the periodicity of changes in xylem structure. Four trees at 770 m were monitored for flowering, branch elongation, leaf litterfall, and xylem cell growth and differentiation in the lower stem, and these events were related to local seasonal patterns of rainfall and temperature. Hurricane Hugo defoliated study trees in September, 1989. Bud-break and branch elongation in March, 1989 were followed by earlywood xylem cell production in the lower stem in April and the onset of flowering in May. Leaf litterfall was greatest between April and June, coinciding with peak branch growth and new leaf formation. Latewood xylem was produced in December. The general phenological pattern was synchronized between trees and over study years. Vessel diameter and density were monitored along with thickness of earlywood and latewood and the former converted to vessel lumen area, a measure of xylem conductance capacity. Annual growth rings were formed with periods of earlywood and latewood production coinciding with traditional summer (rainy) and winter (dry) seasons, respectively, in the Luquillo Mountains. Hurricane defoliation was followed by heavy flowering in 1990, a year of reduced branch elongation and annual xylem ring width, and was associated with maximum vessel lumen area, as was flowering in 1989, prior to the hurricane. Hurricane Hugo provided a perturbation that, through its elicited stress response, allowed for the demonstration of the interplay between flowering, branching, structural growth of xylem, and xylem function.     

祁连山西部青海云杉径向生长对气候因子的响应

利用树木径向生长监测仪(Dendrometer)和自动气象站在祁连山西部对8棵青海云杉的径向生长及环境因子进行了连续监测研究。结果表明: Gompertz函数拟合结果显示,2018—2020年青海云杉的径向生长分别开始于4月19日、4月17日和4月10日,在日平均气温超过5.5 ℃时径向生长开始;青海云杉生长结束时间分别为8月17日、8月21日和7月19日,生长结束时间与生长季末期降水量有关。研究区青海云杉径向生长量受干旱抑制强烈,其中与7月的日均温(负相关)和日降水量(正相关)相关性最高,与生长季初期(5月)的日降水量的相关关系存在年际间差异。     

树木年内径向生长对干旱事件的响应——以贺兰山油松为例

在贺兰山苏峪口国家森林公园,利用径向生长测量仪监测2017和2018年2个生长季内、2个海拔(2010和2330 m)油松的径向生长,研究树木径向生长对干旱的响应。结果表明: 2018年6月的干旱事件使得油松径向生长速率减慢,生长量减小;而7—8月的降水使油松的径向生长重新激活。2018年油松的径向生长主要发生在6—8月,相比2017年延长一个月。油松径向生长与气候因子的响应关系在不同海拔间没有明显的差异。生长季早期干旱对树木径向生长有抑制作用,生长季中后期降水对树木径向生长具有促进作用。该区的气候重建工作中应当充分考虑8月的气候要素对树轮宽度的影响。     

Effect of local heating and cooling on cambial activity and cell differentiation in the stem of Norway spruce (Picea abies)

BACKGROUND AND AIMS The effect of heating and cooling on cambial activity and cell differentiation in part of the stem of Norway spruce (Picea abies) was investigated. METHODS: A heating experiment (23-25 degrees C) was carried out in spring, before normal reactivation of the cambium, and cooling (9-11 degrees C) at the height of cambial activity in summer. The cambium, xylem and phloem were investigated by means of light- and transmission electron microscopy and UV-microspectrophotometry in tissues sampled from living trees. KEY RESULTS: Localized heating for 10 d initiated cambial divisions on the phloem side and after 20 d also on the xylem side. In a control tree, regular cambial activity started after 30 d. In the heat-treated sample, up to 15 earlywood cells undergoing differentiation were found to be present. The response of the cambium to stem cooling was less pronounced, and no anatomical differences were detected between the control and cool-treated samples after 10 or 20 d. After 30 d, latewood started to form in the sample exposed to cooling. In addition, almost no radially expanding tracheids were observed and the cambium consisted of only five layers of cells. Low temperatures reduced cambial activity, as indicated by the decreased proportion of latewood. On the phloem side, no alterations were observed among cool-treated and non-treated samples. CONCLUSIONS: Heating and cooling can influence cambial activity and cell differentiation in Norway spruce. However, at the ultrastructural and topochemical levels, no changes were observed in the pattern of secondary cell-wall formation and lignification or in lignin structure, respectively.     

Earlywood vessels of Castanea sativa record temperature before their formation

The aim of this study was to identify the climatic signal contained in the earlywood vessel size of the ring-porous chestnut (Castanea sativa) and the physiological processes involved in the underlying mechanisms. In order to assign the encoded signal to a specific physiological process, bud phenology and vessel formation were monitored along an elevation transect and chronologies of the size of the first row of earlywood vessels were retrospectively correlated with 40 yr of early spring temperatures. The first vessels appeared in late April to early May, after encoding both a negative temperature signal in February-March (during tree quiescence) and a positive temperature signal in early April (at the time of resumption of shoot growth). We hypothesize that February and March temperatures affect cambial sensitivity to auxin, preconditioning tree responses later in the season. Furthermore, April temperature is related to tree activation whereby new hormone production fosters vessel expansion.     

Concentrations of oxygen and indole-3-acetic acid in cambial region during latewood formation and dormancy development in Picea abies stems

To manipulate the occurrence of latewood formation and cambial dormancy in Picea abies (L.) Karst. stems, potted seedlings were transferred from the natural environment on 9 July, when tracheids early in the transition between earlywood and latewood were being produced, and cultured for up to 5 weeks in a controlled environment chamber having: (1) Warm LD, (25/15C during day/night) and long (16 h) photoperiod, (2) Warm SD, (25/15C) and short (8 h) photoperiod, or (3) Cold SD, (18/8°C) and short (8 h) photoperiod. In Warm LD trees, the radial enlargement of primary-walled derivatives on the xylem side of the cambium, as well as xylem production, continued at the same magnitude throughout the experiment. In Warm SD and Cold SD trees, the radial enlargement of primary-walled derivatives declined and the cambium entered dormancy, both developments occurring faster in the Warm SD trees. The concentrations of indole-3-acetic acid (IAA) was higher in developing xylem tissue than in cambium+phloem tissues, but did not vary with environmental treatment or decrease during the experimental period. The O2 concentration in the cambial region followed the order of Cold SD>Warm SD>Warm LD trees and was <5%, the threshold for the inhibition of IAA-induced proton secretion, for the first 3 weeks in Warm SD and Warm LD trees. Thus, neither latewood formation nor cambial dormancy can be attributed to decreased IAA in the cambial region. Nor does lower O2 concentration in the cambial region appear to be inhibiting the IAA action that is associated with cambial growth.     

Periodicity of cambial activity in Abies balsamea. I. Effects of temperature and photoperiod on cambial dormancy and frost hardiness

The relationship between from hardiness and growth potential, and their dependence on temperature and photoperiod, was investigated in the one-year-old cambium of balsam fir [Abies balsamea (L.) Mill.]. Six-year-old trees were exposed for 9 weeks to either the natural environment or one of 4 controlled environments in the fall (18 September-18 November), spring (12 April–14 June) and summer (19 July – 19 September). The 4 controlled environments were (1) WS, warm temperature (24/20°C in day/night) + short day (8 h). (2) WL. warm temperature (24/20°C) + long day (8 h + 1 h night break), (3) CS. cold temperature (9/5°C) + short day (8 h) and (4) CL, cold temperature (9/5°C) + long day (8 h + 1 h night break). At the beginning and end of each exposure, cambial activity was measured by recording the number of xylem, cambium and phloem cells, frost hardiness was estimated from the cambium's ability to survive freezing to –40°C, and cambial growth potential was deduced from the duration of the cell cycle and the production of xylem, cambium and phloem cells in cuttings cultured for 4 weeks with exogenous indole-3-acetic acid (IAA) under environmental conditions favourable for cambial activity. In the natural environment, frost hardening began in September and was completed in November, while dehardening occurred when the cambium reactivated. CL, CS, and to a lesser extent WS, promoted hardening in the summer and fall, but did not prevent dehardening in the spring. The cambial growth potential in the natural environment declined from a maximum in April to a low level in June, reached a minimum in September, then increased to a high level in November. This potential was promoted by CL and CS on all dates by WL in the summer and fall. The ratio of xylem to phloem induced by IAA treatment was greatest in June and least in September in cuttings from trees exposed to the natural environment, and was increased by CL and CS in the fall. The cambium in intact branches of trees protected from chilling during the fall and winter resumed cell cycling after less than 9 weeks of dormancy, but produced mostly or only phloem in the subsequent growing period. It is concluded that the frost hardiness of the cambium, the IAA-induced cycling of cambial cells, and IAA-induced xylem to phloem ratio vary independently with season, temperature and photoperiod, and that the periodicity of these processes is regulated endogenously.