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。     

水热耦合对沈阳地区油松木质部生长的影响

利用微树芯技术可以从细胞尺度研究树木形成层物候和径向生长的过程,揭示树木生长与气候的关系。油松是我国北方森林的建群树种之一,也是沈阳地区的优势树种。研究2020年整个生长季(4—11月)油松周尺度的形成层及木质部细胞变化,分析油松在沈阳地区的生长规律。结果表明: 油松形成层分裂活动开始于4月初,结束于9月末。木质部从扩大细胞出现(4月中旬)开始生长,到木质化细胞消失(10月下旬)结束,其生长符合“S”型曲线。2020年生长53个/列木质部细胞,最大生长速率(0.55个/列/d)出现在5月末,早晚材细胞于7月末发生转变。在沈阳地区最低温达到0 ℃以上时树木形成层开始活动,影响木质部生长开始和结束的最低临界温度为2～3 ℃。降水在油松整个生长过程中起到促进作用。沈阳地区7月末的高温和水分供给不足是木质部细胞分化形成早晚材的主要因子。     

基于日均温度的华山松径向生长敏感温度研究

树干木质部形成周期与温度密切相关,存在影响形成层活动的敏感温度,但是尚未研究尝试从树木年轮中探索影响径向生长的敏感温度。华山主峰的华山松径向生长对气候变化较敏感,且华山西峰的气象站记录了1953年以来气象资料,这为探索树木径向生长与温度的关系提供了宝贵的材料。以华山主峰的华山松年轮资料和日值温度资料为基础,通过分析历年日平均温度的变化过程和年轮宽度、早材宽度、晚材宽度、最小密度以及最大密度的相关性,尝试探索华山松径向生长与日均温度变化的关系。结果表明春季一定温度的初日时间对华山松径向生长有较大的影响,其中3 ℃和8 ℃初日时间和持续时间对年轮宽度的促进作用最明显,而3 ℃对早材宽度影响较大,8℃初日时间对晚材的影响较大;高于11 ℃的温度会对华山松的径向生长造成限制,其中以11 ℃的作用最明显;而温度的终日时间对年轮特征影响较小。说明3 ℃是早材形成的敏感温度,8 ℃是晚材形成的敏感温度,温度高于11 ℃会对华山松的径向生长构成胁迫。这证明华山松年轮特征中包含了较多的气候信息,形成层活动和木质部的生长存在阈值温度,通过分析不同温度的生长期与年轮特征的关系可以找到影响华山松径向生长的敏感温度。这些敏感的温度是通过什么生理过程影响木质部的形成尚无法得知,但是这为解释树木生长对全球温度升高的响应以及利用华山松年轮重建历史气候提供了重要依据。     

河北塞罕坝樟子松径向生长动态变化及其与气象因子的关系

树木是森林生态系统的基本组成, 其生长受气象因子的影响, 基于此, 该研究通过监测樟子松(Pinus sylvestris var. mongolica)的径向生长, 研究樟子松生长日动态规律、季节动态规律及其与气象因子的关系, 探讨河北塞罕坝地区樟子松森林生态系统对气候变化的响应机制。此外, 以往研究树木生长大多数基于树轮年代学, 缺少短期树木径向生长动态的研究。该研究利用径向生长记录仪监测河北塞罕坝机械林场内樟子松连续3年(2016-2018)的树干径向动态变化。结果表明: 由于树干的水分吸收与蒸腾作用, 樟子松树干径向昼夜变化呈现季节性规律, 可划分为4个阶段: 春季萌动期、夏季生长期、秋冬交替期和冬季休眠期。塞罕坝樟子松树干径向生长开始于每年4月初; 4月初至5月中旬为水分恢复阶段; 5月中旬至7月中旬为快速生长阶段; 7月中旬至10月中旬为缓慢生长阶段; 10月中、下旬生长趋于停止, 并有树干径向收缩现象。以一天为时间尺度, 在快速生长阶段(5月初至7月中旬)樟子松径向生长主要受空气温度的影响; 缓慢生长阶段(7月中旬至10月下旬)降水量、空气温度均影响樟子松径向生长。以15天为时间尺度, 温度对樟子松径向生长的影响显著。结果显示樟子松的生长动态规律及其影响因子, 为未来樟子松生理研究提供参考时间节点, 同时在极端低温与干旱的情况下, 为半干旱地区樟子松的生长状态提供参考依据。     

河北塞罕坝樟子松径向生长动态变化及其与气象因子的关系

树木是森林生态系统的基本组成,其生长受气象因子的影响,基于此,该研究通过监测樟子松(Pinussylvestrisvar.mongolica)的径向生长,研究樟子松生长日动态规律、季节动态规律及其与气象因子的关系,探讨河北塞罕坝地区樟子松森林生态系统对气候变化的响应机制。此外,以往研究树木生长大多数基于树轮年代学,缺少短期树木径向生长动态的研究。该研究利用径向生长记录仪监测河北塞罕坝机械林场内樟子松连续3年(2016–2018)的树干径向动态变化。结果表明:由于树干的水分吸收与蒸腾作用,樟子松树干径向昼夜变化呈现季节性规律,可划分为4个阶段:春季萌动期、夏季生长期、秋冬交替期和冬季休眠期。塞罕坝樟子松树干径向生长开始于每年4月初; 4月初至5月中旬为水分恢复阶段; 5月中旬至7月中旬为快速生长阶段; 7月中旬至10月中旬为缓慢生长阶段; 10月中、下旬生长趋于停止,并有树干径向收缩现象。以一天为时间尺度,在快速生长阶段(5月初至7月中旬)樟子松径向生长主要受空气温度的影响;缓慢生长阶段(7月中旬至10月下旬)降水量、空气温度均影响樟子松径向生长。以15天为时间尺度,温度对樟子松径向生长的影响显著。结果显示樟子松的生长动态规律及其影响因子,为未来樟子松生理研究提供参考时间节点,同时在极端低温与干旱的情况下,为半干旱地区樟子松的生长状态提供参考依据。     

祁连山东部青杄年内径向生长动态对气候的响应

树木径向生长对气候因子的响应是树轮气候学的基础。在我国西北地区,虽然已有大量的树轮-气候响应研究,但是响应分析多基于数理统计结果,缺乏对树木生长过程的理解。基于此,于2013年到2015年在祁连山东部吐鲁沟国家森林公园内通过Dendrometer连续监测了树轮气候重建中常用树种青杄的径向生长,通过平均值法提取2013到2015年生长季内不同时间尺度(1天、7天、10天)的平均径向生长量及对应时间段的平均气候状况。不同时间尺度径向生长量和气候因子的相关分析结果表明,时间尺度的延长削弱了树干水分昼夜变化的干扰,但水分仍然是青杄径向生长的限制因子。为了进一步分析青杄径向生长与气候关系随时间的变化情况,以31天为窗口将日径向生长量与气候要素每隔一天进行滑动相关,结果显示:5、7月青杄的径向生长与降水的关系稳定,都呈显著正相关,但是6月降水的年际变率较大,导致树木可利用水分的变幅也较大,因而树木径向生长与6月水分的响应关系不稳定,存在较大的年际差异,而这可能是一些树轮-气候响应研究中轮宽与6月降水关系不显著的原因。     

昌岭山油松年内径向生长特征及其对气候的响应

利用树木径向生长记录仪对甘肃昌岭山油松径向生长进行了连续三年动态监测,通过平均值法提取了油松净径向生长数据,利用Gompertz函数对油松径向生长曲线进行拟合,分析了油松年内径向生长特征,并结合气象资料分析了油松径向生长对温度和降水的响应,探讨了油松径向生长开始时间与温度的关系以及气候影响油松径向生长的机制。结果表明:(1)油松年内径向生长速率先增大后减小,最大值出现5月下旬到6月上旬,主要生长时间集中在5—8月。两棵油松径向生长开始时间较一致,出现在4月上旬和中旬,但结束时间存在较大差异。(2)生长期油松径向生长与平均气温、最高气温和最低气温均呈负相关,与降水呈正相关,但相关性强弱存在年际差异。(3)昌岭山地区油松径向生长开始的平均气温阈值在7℃左右。生长季温度会通过影响水分条件间接对油松径向生长造成影响,异常充沛的降水可能是2019年6月两棵油松净径向生长量偏多的主要原因。     

阿尔泰山西伯利亚落叶松径向生长对气候变化的分异响应

在全球气候变暖的背景下, 北半球中高纬度地区出现了树轮径向生长对气候变化的分异响应现象, 但是阿尔泰山优势针叶树种对气候因子响应的稳定性还存在不确定性。该研究选择阿尔泰山中段高海拔西伯利亚落叶松(Larix sibirica)样本建立了树轮宽度年表, 并对年表特征及树木径向生长-气候的动态关系进行了分析。结果表明: 生长季初期和中期的气温是研究区树木生长的主控气候因子; 树木径向生长与当年4月的气温显著负相关, 与当年6-7月的气温显著正相关; 研究区西伯利亚落叶松径向生长与当年4月和6-7月的气温发生了分异现象, 表现为随着气候变化, 树木径向生长对生长季初期由高温引起的干旱的响应敏感性越来越强, 而对生长季中期气温的敏感性表现出先减弱再增强的趋势。阿尔泰山西伯利亚落叶松径向生长对气候变化的响应比较敏感, 适合开展树木生长-气候变化的研究; 检验树木径向生长对气候变化分异响应为该区域基于树木年轮开展历史气候重建和提高未来森林生态系统发展趋势预测的准确性提供了科学依据。     

东北南部蒙古栎径向生长对气候变化的响应——以千山为例

蒙古栎是东北森林中最重要的阔叶树种之一.本研究利用树木年代学方法研究中国东北南部千山地区蒙古栎的径向变化,结合1951—2010年的温度和降水等气象数据,利用相关函数分析了树木生长与气候变化的关系,揭示蒙古栎径向生长对气候响应规律.结果表明：研究区4—7月的降水量与蒙古栎年轮宽度呈显著正相关,是限制该地区蒙古栎径向生长的主要限制因子;5月极端最高温度与蒙古栎年轮宽度呈显著负相关,也是影响蒙古栎生长的关键因素.研究期间,蒙古栎年轮宽度与4月降水量的相关显著且稳定,自20世纪80年代开始蒙古栎径向生长对夏季温度的响应敏感性逐渐减弱,对温度的响应表现出从响应夏季温度向响应春季温度的转变.     

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。     

Moisture‐driven xylogenesis in Pinus ponderosa from a Mojave Desert mountain reveals high phenological plasticity

Future seasonal dynamics of wood formation in hyperarid environments are still unclear. Although temperature‐driven extension of the growing season and increased forest productivity are expected for boreal and temperate biomes under global warming, a similar trend remains questionable in water‐limited regions. We monitored cambial activity in a montane stand of ponderosa pine (Pinus ponderosa) from the Mojave Desert for 2 consecutive years (2015–2016) showing opposite‐sign anomalies between warm‐ and cold‐season precipitation. After the wet winter/spring of 2016, xylogenesis started 2 months earlier compared to 2015, characterized by abundant monsoonal (July–August) rainfall and hyperarid spring. Tree size did not influence the onset and ending of wood formation, highlighting a predominant climatic control over xylem phenological processes. Moisture conditions in the previous month, in particular soil water content and dew point, were the main drivers of cambial phenology. Latewood formation started roughly at the same time in both years; however, monsoonal precipitation triggered the formation of more false rings and density fluctuations in 2015. Because of uncertainties in future precipitation patterns simulated by global change models for the Southwestern United States, the dependency of P. ponderosa on seasonal moisture implies a greater conservation challenge than for species that respond mostly to temperature conditions.     

Adjustment Capacity of Maritime Pine Cambial Activity in Drought-Prone Environments

Intra-annual density fluctuations (IADFs) are anatomical features formed in response to changes in the environmental conditions within the growing season. These anatomical features are commonly observed in Mediterranean pines, being more frequent in younger and wider tree rings. However, the process behind IADF formation is still unknown. Weekly monitoring of cambial activity and wood formation would fill this void. Although studies describing cambial activity and wood formation have become frequent, this knowledge is still fragmentary in the Mediterranean region. Here we present data from the monitoring of cambial activity and wood formation in two diameter classes of maritime pine (Pinus pinaster Ait.), over two years, in order to test: (i) whether the differences in stem diameter in an even-aged stand were due to timings and/or rates of xylogenesis; (ii) if IADFs were more common in large trees; and (iii) if their formation is triggered by cambial resumption after the summer drought. Larger trees showed higher rates of cell production and longer growing seasons, due to an earlier start and later end of xylogenesis. When a drier winter occurs, larger trees were more affected, probably limiting xylogenesis in the summer months. In both diameter classes a latewood IADF was formed in 2012 in response to late-September precipitation, confirming that the timing of the precipitation event after the summer drought is crucial in determining the resumption of cambial activity and whether or not an IADF is formed. It was the first time that the formation of a latewood IADF was monitored at a weekly time scale in maritime pine. The capacity of maritime pine to adjust cambial activity to the current environmental conditions represents a valuable strategy under the future climate change conditions.     

Is precipitation a trigger for the onset of xylogenesis in Juniperus przewalskii on the north-eastern Tibetan Plateau?

Background and Aims A series of studies have shown that temperature triggers the onset of xylogenesis of trees after winter dormancy. However, little is known about whether and how moisture availability influences xylogenesis in spring in drought-prone areas.Methods Xylogenesis was monitored in five mature Qilian junipers (Juniperus przewalskii) by microcore sampling from 2009 to 2011 in a semi-arid area of the north-eastern Tibetan Plateau. A simple physical model of xylem cell production was developed and its sensitivity was analysed. The relationship between climate and growth was then evaluated, using weekly wood production data and climatic data from the study site.Key Results Delayed onset of xylogenesis in 2010 corresponded to a negative standardized precipitation evapotranspiration index (SPEI) value and a continuous period without rainfall in early May. The main period of wood formation was in June and July, and drier conditions from May to July led to a smaller number of xylem cells. Dry conditions in July could cause early cessation of xylem differentiation. The final number of xylem cells was mainly determined by the average production rate rather than the duration of new cell production. Xylem growth showed a positive and significant response to precipitation, but not to temperature.Conclusions Precipitation in late spring and summer can play a critical role in the onset of xylogenesis and xylem cell production. The delay in the initiation of xylogenesis under extremely dry conditions seems to be a stress-avoidance strategy against hydraulic failure. These findings could thus demonstrate an evolutionary adaptation of Qilian juniper to the extremely dry conditions of the north-eastern Tibetan Plateau.     

Seasonal Rhythms of Structure and Behaviour of Vascular Cambium in Ficus rumphii

Cambial structure and activity of Ficus rumphii Blume vary withthe changes in local climate. The cambial cells start swellingearly in April prior to the onset of periclinal divisions whichare most frequent in August. Cell division stops in October.During the growth season, initiation as well as cessation ofthe phloem production precedes that of xylem. A moderately hightemperature is correlated with the cambial reactivation. Onceinitiated, the activity continues at relatively low temperatures.Hot and dry environment favours the phloem production, whereashot and moderately humid conditions induce xylogenesis. Thesize and relative proportion of cambial initials also changewith season. Fusiform initials are shorter and broader duringthe rainy season (July–September) than for the rest ofthe year. Multiseriate and triseriate rays, as also the tallrays, outnumber the other types of rays throughout the year. Ficus rumphii, vascular cambium, phenology, climatic variation     

Evidence of threshold temperatures for xylogenesis in conifers at high altitudes

Temperature is the most important factor affecting growth at high altitudes. As trees use much of the allocated carbon gained from photosynthesis to produce branches and stems, information on the timing and dynamics of secondary wood growth is crucial to assessing temperature thresholds for xylogenesis. We have carried out histological analyses to determine cambial activity and xylem cell differentiation in conifers growing at the treeline on the eastern Alps in two sites during 2002–2004 with the aim of linking the growth process with temperature and, consequently, of defining thresholds for xylogenesis. Cambial activity occurred from May to July–August and cell differentiation from May–June to September–October. The earliest start of radial enlargement was observed in stone pine in mid-May, while Norway spruce was the last species to begin tracheid differentiation. The duration of wood formation varied from 90 to 137 days, depending on year and site, with no difference between species. Longer durations were observed in trees on the south-facing site because of the earlier onset and later ending of cell production and differentiation. The threshold temperatures at which xylogenesis had a 0.5 probability of being active were calculated by logistic regressions. Xylogenesis was active when the mean daily air temperature was 5.6–8.5°C and mean stem temperature was 7.2–9°C. The similar thresholds among all trees suggested the existence of thermal limits in wood formation that correspond with temperatures of 6–8°C that are supposed to limit growth at the treeline. Different soil temperature thresholds between sites indicated that soil temperature may not be the main factor limiting xylogenesis. This study represents the first attempt to define a threshold through comparative assessment of xylem growth and tissue temperatures in stem meristems at high altitudes.     

Duration of xylogenesis in black spruce lengthened between 1950 and 2010

Background and Aims

Reconstructions have identified the 20th century as being uniquely warm in the last 1000 years. Changes in the phenology of primary meristems converged toward increases in length of the growing season. Has the phenology of secondary meristem changed during the last century, and to what extent?

Methods

Timings of wood formation in black spruce, Picea mariana, were monitored for 9 years on a weekly timescale at four sites in the boreal forest of Quebec, Canada. Models for assessing xylem phenology were defined and applied to reconstruct onset, ending and duration of xylogenesis between 1950 and 2010 using thermal thresholds on chronologies of maximum and minimum temperatures.

Key Results

All sites exhibited increasing trends of both annual and May–September temperatures, with the greatest changes observed at the higher latitudes. Phenological events in spring were more affected than those occurring in autumn, with cambial resumptions occurring 0·5–0·8 d decade⁻¹ earlier. The duration of xylogenesis has lengthened significantly since 1950, although the models supplied wide ranges of variations, between 0·07 and 1·5 d decade⁻¹, respectively.

Conclusions

The estimated changes in past cambial phenology demonstrated the marked effects of the recent increase in temperature on the phenological traits of secondary meristems. In the long run, the advancement of cambial activity could modify the short time window for growth of boreal species and dramatically affect the dynamics and productivity of trees in these temperature-limited ecosystems.     

Variations in the timing of different phenological stages of cambial activity in Abies pindrow (Royle) along an elevation gradient in the north-western Himalaya

High mountains around the globe are some of the most vulnerable ecosystems to climate change and of great concern for conservation. The Himalayan Mountains are experiencing a higher warming than average global warming, which can significantly impact their biodiversity, vegetation distribution and ecosystem structure. There is a need to study the process of wood formation in Himalayan conifers to have a better understanding of their growth responses to predicted climate change. Variations in the climatic factors regulating cambial activity would result in changes in the timing of cambial phenology. In this study, the variations in the timing of different stages of cambial phenology (cell enlargement stage, wall-thickening stage and cell maturation stages) in pindrow fir (Abies pindrow) were investigated from anatomical observations of wood microcores collected during 2014-15 along an elevation range of c.2300−3000 m asl in the north-western Himalaya. The onset of all three cambial phenological stages was significantly correlated with elevation, with onset of cambial activity happening more than a week earlier at the lowest elevation than at the highest elevation site. Although the termination of wall-thickening and maturation stage appeared minimally related to elevation, the cell-enlargement stage showed significant correlation with elevation, with tracheid formation ceasing approximately three weeks earlier in trees at the highest elevation. The timing of these phenological stages did not show strong variations between the two study years. Our findings provide new data on the timings of cambial phenophases and help to understand tree growth response to ongoing changing climate in the Himalayan region.     

Hormonal signals involved in the regulation of cambial activity, xylogenesis and vessel patterning in trees

The radial growth of plant stem is based on the development of cribro-vascular cambium tissues. It affects the transport efficiency of water, mineral nutrients and photoassimilates and, ultimately, also plant height. The rate of cambial cell divisions for the assembly of new xylem and phloem tissue primordia and the rate of differentiation of the primordia into mature tissues determine the amount of biomass produced and, in the case of woody species, the wood quality. These complex physiological processes proceed at a rate which depends on several factors, acting at various levels: growth regulators, resource availability and environmental factors. Several hormonal signals and, more recently, further regulatory molecules, have been shown to be involved in the induction and maintenance of cambium and the formation of secondary vascular tissues. The control of xylem cell patterning is of particular interest, because it determines the diameter of xylem vessels, which is central to the efficiency of water and nutrient transport from roots to leaves through the stem and may strongly influence the growth in height of the tree. Increasing scientific evidence have proved the role of other hormones in cambial cell activities and the study of the hormonal signals and their crosstalking in cambial cells may foster our understanding of the dynamics of xylogenesis and of the mechanism of vessel size control along the stem. In this article, the role of the hormonal signals involved in the control of cambium and xylem development in trees and their crosstalking are reviewed.